(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Bulletin of the British Museum (Natural History) Entomology"




fh- 



\i\i 



BULLETIN OF 

THE BRITISH MUSEUM 

(NATURAL HISTORY) 



ENTOMOLOGY 
Vol. XXVII 



BRITISH MUSEUM (NATURAL HISTORY) 
LONDON : 1973 



7 DEC 3973 



Printed in England by Unwin Brothers Limited 



( 



-7DECJ973 

i 



CONTENTS 

ENTOMOLOGY VOLUME XXVII 

No. I. A revision of the genus Masalia (Lepidoptera: Heliothidinae). By 

P. R. Seymour i 

No. 2. A taxonomic review of the species of Cinara Curtis occurring in 

Britain (Hemiptera: Aphididae) . By V. F. Eastop ioi 

No. 3. The Simuliidae described by N. Baranov and their types (Diptera). 

By R. W. Crosskey & B. V. Peterson in- 

No. 4. Revisional notes on African Charaxes (Lepidoptera: Nymphalida 

Part VIII. By V. G. L. van Somiki \ 215 

No. 5. On European Pteromalidae (Hymenoptera): a revision of Cleonytnus, 
Eunotus and Spaniopus, with descriptions of new genera and species. 
By Z. Boucek 265 

No. 6. The ant genera of Wesl Africa: a synonymic synopsis with keys 

(Hymenoptera: Formicidae). By B. Bolton 317 

No. 7. Contributions towards a revision of Myrsidea Waterston. VII. 

(Phthiraptera: Amblycera: Menoponidae) . By B. K. Tan dan 369 

No. 8. A revision of the Lecanodiaspis Targioni-Tozzetti (Homoptera: 

Coccoidea) of the Ethiopian Region. By C. J. Hodgs< >\ 411 

Index to Volume XXVII 453 



REVISION OF THE GENUS 






- 



A 

MAS AH A (LEPIDOPTERA : 

HELIOTHIDINAE) 



P. R. SEYMOUR 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 27 No. 1 

LONDON : 1972 



A REVISION OF THE GENUS MASALIA 
(LEPIDOPTERA : HELIOTHIDINAE) 




BY 



PAUL ROY SEYMOUR 






Pp. i-ioo; 10 Plates, 113 Text-figures, 11 Maps 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 27 No. 1 

LONDON : 1972 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 27 No. 1 of the Entomolog- 
ical series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Ent. 



Trustees of the British Museum (Natural History) 1972 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



Issued 3 March, 1972 



A REVISION OF THE GENUS MASALIA 
(LEPIDOPTERA : HELIOTHIDINAE) 

By P. R. SEYMOUR 



CONTENTS 

Synopsis 

Material Studied . 
Acknowledgements 
Treatment 
Masalia Moore 

Redescription 

Diagnosis . 

Historical survey 

Generic affinities and Distribution 

Groups within the genus 

Key to the species and subspecies 

Descriptions of the species and subspecies 
References ...... 

Index ....... 

SYNOPSIS 



Page 

3 

3 

4 

4 

6 

6 

8 

8 

io 

io 

ii 

19 

97 

98 



The genus Masalia Moore is recalled from synonymy and fully revised. A key is given to 
the 38 species (three new) and 31 subspecies (five new) recognized as valid. Seventeen specific 
and two subspecific synonyms are newly established. 



MATERIAL STUDIED 

The large collection of Heliothidine moths in the British Museum (Natural History) 
formed the nucleus of the material used for this revision. Type and other important 
material was borrowed from collections in the following museums : 

Mus£e Royal de l'Afrique Centrale, Tervuren (MRAC, Tervuren) ; Museo Civico 
di Storia Naturale, Genoa (MCSN, Genoa) ; Museum Alexander Koenig, Bonn 
(MAK, Bonn); Museum National d'Histoire Naturelle, Paris (MNHN, Paris); 
Naturhistoriska Riksmuseet, Stockholm (NR, Stockholm) ; University Museum, 
Oxford (UM, Oxford) ; Museum fur Naturkunde der Humboldt-Universitat, Berlin 
(MNHU, Berlin); and Zoologisches Sammlung des Bayerischen Staates, Munich 
(ZSBS, Munich). 

The abbreviations given in brackets are those used throughout the text in listing 
the material examined. 

Specimens were examined of all the species placed in Timora Walker, 1856, the 
genus from which Masalia has been extracted and with which it was previously 



4 P. R. SEYMOUR 

synonymized. Other species within the subfamily with affinities to Timora were 
also investigated. Except for four, all the holo- or lectotypes of valid names and 
synonyms now included in Masalia have been seen. The four exceptions were 
M. epimethea (Viette), M. prochaskai (Viette), M. leucosticta vinida (Berio) and 
M. lancea (Berio) ; for each of these an author-verified specimen was examined and 
paratype photographs were seen of vinula and lancea. 

ACKNOWLEDGEMENTS 

I wish to thank the following specialists for their kindness in lending type and 
other material: Mons. L. A. Berger, Belgium; Dr E. Berio, Italy; Dr J. Bourgogne, 
France; Dr H. J. Hannemann, Germany; Mr E. Taylor, England; and Dr E. Todd, 
U.S.A. 

I am also most grateful to Dr W. Forster, Germany; Dr G. Hallin, Sweden; and 
Dr P. Viette, France, who, in addition to lending material, have given help with 
this paper in many other ways. 

TREATMENT 

The layout of the accounts of the species and subspecies has been standardized 
and arranged in the following sequence: fore and hind wing, genitalia, material 
examined, distribution, and remarks (diagnosis and comment). Mention is made 
of the antennae, proboscis quotient and 8th abdominal tergum only when these 
differ from the norm or, in the case of the antennae, when they are sexually di- 
morphic. The 'proboscis quotient' is the length of the proboscis divided by the 
length of the long axis of the eye. 

The presence or absence of an areole in the fore wing and the fore wing length are 
stated; the latter is measured from the fore wing apex to the centre of the meso- 
thorax. The measurements given are in millimetres and are for the smallest and 
largest specimen of each sex; the number of specimens checked is given in paren- 
theses. Holotype and lectotype measurements have also been included in order to 
give a standard set of figures, for in a number of the earlier descriptions the points 
between which measurements were taken was not stated. 

Three fore wing regions are recognized: costal, central and anal (shown against 
fore wing veins in Text-fig. i). In the descriptions of the pattern, 'upper central 
longitudinal streak' refers to the streak passing through the radial half of the cell 
and beyond between Mi and Mi (Text-fig. 2) ; 'lower central longitudinal streak' 
refers to the streak passing through the cubital half of the cell and beyond between 
M 2 and Mz (Text-fig. 3) ; 'anal longitudinal streak' refers to the streak following 
the anal fold (Text-fig. 2). The colour terms used refer to the general names given 
in the colour diagrams of Methuen's 1967 Handbook of Colour. 

The fore and hind wing upper surface of each species and subspecies has been 
illustrated by a photograph. Illustrations of variants to indicate the range of 
variation of wing-pattern found are also included (sex-linked variation is indicated) . 
For the illustrations fresh, well marked specimens were given preference over faded 
or worn types. 



REVISION OF GENUS MASALIA 



costal region 



central region 




anal region 



anal longitudinal streak 



upper central 
longitudinal streak 




lower central 
longitudinal streak 



Figs 1-4. Fore wing longitudinal markings of Masalia and their terms. 4, forked 
lower central longitudinal streak. 



6 P. R. SEYMOUR 

The male genitalia is illustrated by figures of the scobinate bar and proximal end 
of the vesica (Text-fig. 14). Differences centre on bar-shape and on the size and 
number of spicules. Differences occur both inter- and intraspecifically, and between 
a number of species there is overlapping variation. Although the diagnostic value 
of these characters is thereby reduced they are nevertheless particularly important 
in aiding identification of male specimens, since in the key female characters are 
often used. Figures of the scobinate bar of all species are included to provide a 
comparative set. Extremes have been chosen for species having a wide degree of 
bar variation, whilst for others a typical representative specimen has been used. 
For the female, figures are given showing the lateral view of the papilla analis of the 
species and subspecies in which the papilla analis is modified in form. 

The data of specimens examined are listed. The information has been taken 
from their attached labels and, in the case of type-material, from the original 
descriptions where additional information is recorded. The locality data of type- 
material is stated as given on the specimen, but changes in country or regional name 
and in spelling (where traced) have been inserted in square brackets. The changes 
and alteration to spelling are based on names in the 1965 Times Index-Gazetteer of 
the World and on current usage in African countries whose names have since changed. 
Other material has been listed geographically to country (using the current 
country-names) and alphabetically to locality. Unless otherwise stated specimens 
are in the collection of the British Museum (Natural History). 

MASALIA Moore gen. rev. 

Masalia Moore, 1881: 364. Type-species: Masalia radiata Moore, by original designation. 
Pradatta Moore, 1881 : 364. Type-species: Pradatta beatrix Moore, by original designation. 

[Synonymized with Timora Walker by Hampson, 1903 : 103.] 
Curubasa Moore, 1881 : 366. Type-species: Alaria lanceolata Walker, by original designation. 

[Synonymized with Timora Walker by Hampson, 1903 : 103.] 
[Timora Walker; Hampson, 1903 : 103. Masalia Moore synonymized with Timora Walker.] 
Lecerfia Dumont, 1920 : 102. Type-species: Lecerfia chitinipyga Dumont, by monotypy. 

[Synonymized with Timora Walker by Draudt, 1935 : 197.] 

Redescription. Head. Clypeus differentiated into a bulbous plate extending well forward 
from the eyes and a ventral upcurved semicircular plate with a dorsal protruding lip (Text-fig. 
8), the bulbous plate clothed in short adpressed, hair-like scales, and the ventral plate glabrous. 
Proboscis short, proboscis quotient from 2, but only exceptionally exceeding 3; distally bearing 
numerous well developed sensory papillae. Antenna with 50 to 60 flagellar segments. 

Thorax. Prothoracic tibia with an epiphysis and a terminal pair of large modified spines, 
of which the inner is the shorter (Text-fig. 6), or occasionally with a single modified spine; 
otherwise without spines. Mesothoracic tibia with a terminal pair of subequal spurs and from 
1 to 6 spines distad (Text-fig. 5). Metathoracic tibia with a terminal and medial pair of sub- 
equal spurs and from 1 to 6 spines distad, spines restricted to below level of medial spurs (Text- 
fig- 7)- 

Wings. Fore wing venation with Sc, Ri, Mi, Mi, M%, Cui & , C«n, and \A as in Text-fig. 9. 
■zA usually absent; when present, weakly developed and anastomosing proximally with lA. 
i?2 to i?5 present, arrangement variable. Areole present or absent; when present, between R% 
and i?3+4 or R% and i?3+4+s. 

Hind wing venation with Sc + Ri anastomosing proximally or approximating with Rs. 
M2 absent (Text-fig. 9) . Rs and Mi either divided or stalked from lower angle of cell ; venation 
otherwise constant. 



REVISION OF GENUS MASALIA 




tibia 



spines 



terminal 
spurs 



modified 
spines 



\tarsus 





\ medial 
spurs 




Figs 5-8. 



Structures of Masalia. 5, mesothoracic leg. 6, tibia of prothoracic leg. 
7, metathoracic leg. 8, head capsule, fronto-lateral view. 



8 P. R. SEYMOUR 

Abdomen. Posterior margin of eighth abdominal tergum from straight to centrally emar- 
ginate (Text-figs 20, 21). 

Genitalia. Male with uncus simple, terminally hooked; valve simple, apically slightly dilate, 
corona spiculate (Text-figs 12, 13). Aedeagus with apex obliquely truncate with a scobinate 
bar; vesica membranous, slightly spiral and with a scale-like cornutus (Text-fig. 14). Female 
papilla analis either simple [i.e., membranous and rounded] (Text-figs 15, 16), or modified 
[sclerotized] and of variable shape being rounded, folded, angled or digitate (Text-figs 17-19). 
Ductus bursae elongate, corpus bursae with 1 to 4 signa and an appendix bursae; ductus, 
corpus and appendix bursae membranous, simple or ribbed (Text-figs 15, 18). 

Diagnosis. Vesica in males with a proximal scale-like cornutus. 

Historical survey. Moore (1881) erected the genus Masalia for two new species 
M. radiata and M. irrorata, designating the former as type-species. The second of 
these two species is now assigned to Timora Walker. Moore's paper is descriptive 
but not diagnostically orientated and reflects a disregard of earlier work. 

Masalia was synonymized with Timora by Hampson at the beginning of the 
century and has remained a synonym of it until the present paper. Hampson's 
revision of Timora was important in bringing together a number of naturally related 
species but, unfortunately, the characters chosen to distinguish species of Timora 
from other genera are inadequate. 

Later revisions of Timora by Warren (191 1, 1913) and Gaede (1935) closely 
followed the work of Hampson. The names of species subsequently described in 




2A 



Figs 9-1 1. Wing structures and markings of Masalia. 9, fore and hind wing venation. 
10, M . perstriata $, fore wing venation. 11, transverse wing markings typical of decorala- 
group species. 



REVISION OF GENUS MASALIA 



scobinate bar 




Figs 12-14. Structures of male genitalia of Masalia. 12 and 13, valves. 14, aedeagus 

with everted vesica. 



io P. R. SEYMOUR 

Timora were brought together, but there was much confusion, for few types and 
little material had been examined. Recent relevant papers are concerned with 
descriptions of new species and are widely scattered in the literature. 

Hampson's (1903) synonymy of Masalia with Timora was primarily based on 
prothoracic tibial features, namely the absence of spines except for a single or pair 
of terminal modified spines. On examining Heliothidine material in the collection 
of the British Museum (Natural History), the same type of prothoracic tibia was 
found in nine other genera; the number of species involved was comparatively few 
(29) but sufficient to discount the value of the prothoracic tibial features at the 
generic level. Unfortunately, no satisfactory combination of characters could be 
found to separate Timora sensu Hampson from other related genera. 

Following investigation of the male genitalia, a scale-like cornutus was found at 
the proximal end of the vesica in a number of species assigned to Timora (but 
absent in the type-species, Timora senegalensis (Guenee). In these species, the 
cornutus was present in all the specimens examined and was similar in appearance 
from one species to another and though irregular in form, varied only slightly in 
position relative to the scobinate bar. On the presence of a cornutus and similarity 
of other characters these species are regarded as comprising a genus for which the 
name Masalia is recalled from synonymy. Within the Heliothidinae, the vesical 
armature has been found diagnostic in two other genera, Helicoverpa Hardwick, 
1965, with a helical row of spicules or spicule clusters, and Adisura Moore, 1881, 
with a small number of elongate terminal spicules. 

Generic affinities and distribution. Masalia has affinities with Timora, 
Adisura and Canthylidia Butler, 1886, but differs from them in the presence in the 
male of a scale-like cornutus. As in Masalia the prothoracic tibia in Timora and a 
number of species of Canthylidia has one or a pair of subequal apical modified spines. 
In Masalia and Timora the prothoracic tibia is otherwise unspined. In Canthylidia 
there is often one modified spine, but this may be absent. In Adisura the pro- 
thoracic tibia is devoid of modified spines, but other spines are usually present. In 
all four genera the proboscis is short ; in Masalia and Timora the quotient is usually 
2-3; in Adisura and Canthylidia usually between 3 and 4. 

Masalia is distributed across Africa, the Malagasy Republic, Saudi Arabia, 
southern Iran, West and East Pakistan, India, Ceylon, China, Lombok, Flores, and 
northern Australia. 

Groups within the genus. Although there is much structural uniformity 
within the genus, differences in wing pattern and colouring are quite marked. 
Shared and differing characters are met with in an assortment of combinations. A 
number of species-groups are recognized, but it is not suggested that they are entirely 
natural, for some of the characters used may well be affected by parallel evolution. 

The fissifascia-group. Two species, philbyi and fissifascia, characterized by a 
white, distally forked, lower central longitudinal streak on the fore wing (Text-fig. 4). 
Distribution : East Africa, Saudi Arabia, southern Iran and Afghanistan. 

The decor ata-gxoup. The species are decorata, leucostica, funebris, prochaskai and 
sublimus; fore wing with transverse rather than longitudinal markings and with 



REVISION OF GENUS MASALIA n 

pink (or red) and yellow colouring (Text-fig. n). Distribution: Africa, Malagasy 
Republic, Afghanistan, India and Ceylon. 

The galatheae-group. The species are distincta, cruentata and galatheae (the latter 
two being regarded as a species-complex) ; fore wing with the costal and anal regions 
pink, and central region white to yellowish white. Although their fore wings are 
white, flaviceps and hololeuca are also included. In a few specimens of flaviceps, 
almost imperceptible demarcation between the costal, central and anal regions can 
be traced, revealing the pattern found in galatheae. M. hololeuca and M . flaviceps 
are the only species within the genus having white fore wings. Distribution : Africa, 
India and China. 

The radiata-group. The species are radiata, rubristria, beatrix, epimethea, rosacea, 
roseivena and flavistrigata ; fore wing with the costal and anal regions, and parts of 
central region, reddish brown to brown, with a yellowish white, upper central 
longitudinal streak and usually with an anal streak of the same colour. Distribution : 
Africa, Malagasy Republic, India, Lombok, Flores and northern Australia. 

M. latinigra and M . cheesmanae are regarded as a species complex characterized 
by a single modified spine on the prothoracic tibia. Distribution : Africa. 

M. albida, with its long proboscis (proboscis quotient 6) and distinctive pattern, 
and M. perstriata, also with a long proboscis and node-like distension of the costa 
of the fore wing (Text-fig. 10), are species distinct from each other and from others 
within the genus. 

The remaining species do not fit into convenient groupings. 

Distribution. The genus is predominantly Afro-Indian. Of the 38 species, 15 
are African endemics and 15 endemic to India (one extending northwestward into 
China). A further four species occur both in Africa and India (one also occurring 
in Ceylon). Of the remaining four, three occur in the Malagasy Republic, the 
fourth being found in northern Australia and the islands Lombok and Flores, near 
northern Australia. 



KEY TO THE SPECIES AND SUBSPECIES 

The taxonomic features of the Heliothidinae suggest that a large number of 
species have undergone recent speciation. There is a high degree of structural 
uniformity within the subfamily, and marked superficial variation within species; 
the species are separated by only slight discontinuities. The key has been con- 
structed as far as possible on the basis of non-sexual characters, but the use of the 
papilla analis of the female and, less often, the aedeagus and antennae of the male 
has been unavoidable in some couplets. Because of the limited number of structural 
characters, pattern and colour have been widely used. 

1 Fore wing upper surface with a white to yellowish white upper central longi- 

tudinal streak as in Text-fig. 2 or with a forked lower central longitudinal 
streak as in Text-fig. 4 ......... 2 

Fore wing upper surface without a white upper, or forked lower, central 

longitudinal streak .......... 22 



12 P. R. SEYMOUR 

2 (i) Fore wing upper surface with a white forked lower central longitudinal streak 

(following Cu\ and dividing with it along Mz-Cu\^ ..... 3 

- Fore wing upper surface without a white forked lower central longitudinal 

streak ............. 5 

3 (2) Fore wing upper surface light yellow to brownish orange. (Arabian, Iranian 

or Afghanistan species) ..... M. philbyi (Brandt) (p. 19) 

- Fore wing upper surface greyish red. (East African species) ... 4 

4 (3) Fore wing upper surface with the white forked streak outlined with light brown. 

M. fissifascia fissifascia (Hampson) (p. 23) 

- Fore wing upper surface with the white forked streak not outlined with light 

brown ...... M. fissifascia roseata (Pinhey) (p. 24) 

5 (2) Fore wing with areole .......... 6 

- Fore wing without areole .......... 18 

6 (5) Madagascan, African, Arabian or Iranian species. ..... 7 

- Indian species ............ 13 

7 (6) Madagascan species. 

Fore wing with pointed apex (Text-fig. 23) ; upper surface brownish 
orange, costal margin white .... M. epimethea (Viette) (p. 61) 

- African, Arabian or Iranian species ........ 8 

8 (7) Female genitalia with unmodified papilla analis (membranous and of rounded 

shape) (Text-figs 15, 16) ......... 9 

- Female genitalia with modified papilla analis (sclerotized and of variable shape, 

rounded, folded or digitate) (Text-figs 17, 18, 19) . . . . . 12 

9 (8) Fore wing upper surface with costal region pastel to greyish red; central 

region white, with a dark brown to black, lower central longitudinal streak, 
and a faint to well developed dark brown to black streak between R$ and M\ 
(PI. 6, figs 177, 178) . . . M. beatrix trifasciata (Hampson) (p. 61) 

- Fore wing upper surface with costal region pale yellow to greyish orange . 10 

10 (9) Abdomen with posterior margin of 8th tergum ridged and centrally emarginate 

(Text-fig. 20). 

Fore wing upper surface with a white anal longitudinal streak and with 
faint to well developed white streaks between Mi and M3, between M% and 
Cui a , and between C«i a and Cwib (PI. 10, figs 223, 224). 

M. albida (Hampson) (p. 93) 

- Abdomen with posterior margin of 8th tergum not ridged ; straight or centrally 

only slightly emarginate (Text-fig. 21). . . . . . . 11 

11 (10) Hind wing upper surface light to yellowish brown. 

M. rubristria rhodomelaleuca (Berio) (p. 58) 

- Hind wing upper surface white. 

M. perstriata fuscostriata (Brandt) (part) (p. 96) 

12 (8) Fore wing apex rounded (Text-fig. 22). Female genitalia with surface of 

papilla analis spiculate (striate appearance) ; dorso-laterally not sericate, as 

in Text-fig. 86 . . . . M. terracottoid.es (Rothschild) (p. 75) 

- Fore wing apex pointed (Text-fig. 23) . Female genitalia with surface of papilla 

analis not spiculate but dorso-laterally sericate, as in Text-fig. 72. 

M. ftavistrigata (Hampson) (part) (p. 65) 

13 (7) Australian species. 

Fore wing upper surface with a pink (or pink suffused with reddish brown) 
lower central longitudinal streak ; costal and anal regions pink, central region 
white ........ M. roseivena (Walker) (p. 63) 

Indian species ............ 14 

14 (13) Female genitalia with unmodified papilla analis (membranous and of rounded 

shape) (Text-figs 15, 16) ......... 15 



REVISION OF GENUS MASALIA 



«3 




Figs 15-19. Structures of female genitalia of Masalia. 15 and 16, papilla analis 
simple (membranous and rounded). 17-19, papilla analis modified (sclerotized). 



i 4 P. R. SEYMOUR 

- Female genitalia with modified papilla analis (sclerotized and of variable shape, 

rounded, folded, or digitate) (Text-figs 17, 18, 19) ..... 16 

15 (14) Female genitalia with papilla analis of right-angle triangular shape (Text-fig. 

93). Fore wing upper surface light greyish or brownish orange; marginal 
cilia with banded appearance, proximally brownish orange, distally white, 
colour separated by a well defined line as in PI. 8, figs 208, 209. 

M. tost a Moore (part) (p. 80) 

- Female genitalia with papilla analis not of right-angle triangular shape. Fore 

wing upper surface with costal, central and anal regions light orange to 
pastel-red or with central region white with light red to pastel-red streaks. 

M. beatrix beatrix (Moore) (part) (p. 59) 

16 (14) Fore wing upper surface pinkish white. Female genitalia with terminal spines 

on the papilla analis (Text-fig. 19). 

Hind wing upper surface reddish golden brownish orange. 

M. rosacea Hampson (p. 62) 

- Fore wing upper surface pale yellow, light to brownish orange or light brown. 

Female genitalia without terminal spines on the papilla analis (Text-fig. 18) 17 

17 (16) Hind wing upper surface either white or greyish to brownish orange. 

M. radiata terracotta Hampson (part) (p. 52) 

- Hind wing upper surface brown . . . M. radiata radiata Moore (p. 52) 

18 (5) African species ........... 19 

- Indian species. 

Fore wing upper surface pale yellow or light to brownish orange. Hind 
wing upper surface white or greyish to brownish orange. 

M. radiata terracotta Hampson (part) (p. 52) 

19 (18) Female genitalia with unmodified papilla analis (membranous and of rounded 

shape) (Text-figs 15, 16) ......... 20 

- Female genitalia with modified papilla analis (sclerotized and of variable shape, 

rounded, folded or digitate) (Text-figs 17, 18, 19) . . . .21 

20 (19) Fore wing upper surface with greyish rose, light brown, brown or reddish 

brown markings. (West Africa) M. rubristria rubristria (Hampson) (p. 54) 

- Fore wing upper surface with pale to pastel or greyish red markings. (Central, 

East and southern Africa) . M. rubristria transvaalica (Distant) (p. 57) 

21 (19) Male genitalia with cornutus . M. bimaculata cornia subsp. n. (part) (p. 90) 

- Male genitalia without cornutus. 

M. bimaculata pluritelifora (Berio) (part) (p. 90) 

22 (1) Prothoracic tibia with a single terminal modified spine (Text-fig. 74) . . 23 

- Prothoracic tibia with a pair of terminal modified spines (Text-fig. 6) . 27 

23 (22) Fore wing with rounded apex (Text-fig. 22). Female genitalia with un- 

modified papilla analis (membranous and of rounded shape) (Text-figs 15, 16) 24 

- Fore wing with pointed apex (Text-fig. 23). Female genitalia with modified 

papilla analis (sclerotized and of variable shape, rounded, folded or digitate) 
(Text-figs 17, 18, 19) . . . M. flavistrigata (Hampson) (part) (p. 65) 

24 (23) Fore wing upper surface with a distinct broad, dark brown to black lower 

central longitudinal streak . . M. latinigra latinigra (Hampson) (p. 68) 

- Fore wing upper surface with at most an indistinct narrow lower central 

longitudinal streak. .......... 25 

25 (24) Hind wing with upper surface white to orange-white. 

M. cheesmanae cheesmanae subsp. n. (p. 69) 

- Hind wing upper surface brownish orange to brown ..... 26 

26 (25) Male antenna with simple flagellar segments (Text-fig. 79). (Known only from 

southern Sudan) . . . M. cheesmanae tamburensis subsp. n. (p. 70) 

- Male antenna with lamellate flagellar segments (Text-fig. 77). (Known only 

from Ethiopia) .... M. latinigra dangilensis subsp. n. (p. 69) 



REVISION OF GENUS MASALIA 



15 





21 






Figs 20-24. Structures of Masalia. 20 and 21. 8th abdominal terga, dorsal view. 
22-24, fore wing apices rounded (22) and pointed (23) and (24). 



16 P. R. SEYMOUR 

27 (22) Fore wing upper surface with a series of white postmedial dots or dashes (Text- 

figs 32, 33) . 28 

- Fore wing upper surface without a series of white postmedial dots or dashes . 36 

28 (27) Fore wing upper surface from light to greyish yellow with distinct pink to red 

markings ............ 29 

- Fore wing upper surface not of this colour and pattern ; if the colour of the fore 

wing upper surface is yellow and greyish red (variety of M. quilengesi), the 
yellow merges almost imperceptibly into the greyish red. .... 35 

29 (28) Fore wing upper surface with the proximal half of the costal region pink to red 

as in PI. 2, fig. 129, or light to greyish yellow, traversed by a pink to red 
longitudinal streak as in PI. 2, fig. 127 ....... 30 

- Fore wing upper surface with the proximal half of the costal region light to 

greyish yellow, not traversed by a pink to red longitudinal streak as in PI. 1, 

fig- 124 33 

30 (29) African species 

- Madagascan species. 

Fore wing upper surface with a wide pink longitudinal band occupying 
nearly the whole costal region (PI. 2, fig. 132) M. prochaskai (Viette) (part.) (p. 34) 

31 (30) Fore wing upper surface with the post- and antemedial pink markings con- 

tiguous, as in PI. 2, figs 129, 130 ........ 32 

- Fore wing upper surface with the post- and antemedial pink markings not 

contiguous, as in PI. 2, fig. 127 M. leucosticta leucosticta (Hampson) (p. 30) 

32 (31) From northern Tanzania, Kenya, Ethiopia or Somalia. Fore wing usually with 

areole ....... M. leucosticta vinula (Berio) (p. 32) 

- From southern Tanzania or southern Congo (Kinshasa). Fore wing without 

areole. ...... M. leucosticta joiceyi (Prout) (p. 33) 

33 (29) From Africa or Arabia .......... 34 

- From India. 

Male genitalia with scobinate bar and cornutus (Text-fig. 34). 

M. decorata decorata (Moore) (p. 25) 

34 (33) Fore wing upper surface with post medial dots arranged more or less centrally 

within the pink to red postmedial band (Text-fig. 32). 

M. decorata metarhoda (Druce) (p. 27) 

- Fore wing upper surface with postmedial dots arranged along the inner margin 

of the pink to red postmedial band (Text-fig. 33). 

M. decorata albiseriata (Druce) (p. 27) 

35 (28) Fore wing upper surface with the costal and central regions bluish red, anal 

region brown. Hind wing upper surface brown . M. funebris (Berio) (p. 33) 

- Fore wing upper surface light yellow suffused with pastel-red. Hind wing 

upper surface pale yellow .... M. quilengesi sp. n. (part) (p. 74) 
Fore wing upper surface with white orbicular and discocellular dots (Text- 
fig- 25) 37 

Fore wing upper surface without white orbicular and discocellular dots . . 39 

Fore wing upper surface with anal region pink ...... 38 

Fore wing upper surface with anal region yellowish white. 

M. disticta albirosea (de Joannis) (p. 38) 
Fore wing upper surface with central region yellowish white (PI. 2, fig. 136). 

M. disticta disticta (Hampson) (p. 36) 
Fore wing upper surface with central region pink, apart from a pale yellow basal 

dash (PL 2, fig. 137) . . . M. disticta flavirosea (Hampson) (p. 39) 

Fore wing upper surface white or with costal and anal regions yellowish white 

and central region white ......... 40 

Fore wing upper surface not as above . . . . . . .41 

Fore wing with areole. (Indian species) . . M. hololeuca (Hampson) (p. 50) 



36 (27) 


37 


(36) 


38 (37) 


39 


(36) 


40 


(39) 



REVISION OF GENUS MASALIA 



17 



Fore wing without areole. (African species) . M. flaviceps (Hampson) 

Fore wing upper surface with costal region white, white with margin pink, pale 
to light yellow, or pink to dull red; central region white or pale to light 
yellow; anal region light yellow or pink to dull red. Costal and anal regions, 
or anal region only, of darker colour than central region ; otherwise immacu- 
late (PI. 3, figs 138 to 147 and PI. 4, figs 150 to 152) . . . . 

Fore wing upper surface not as above ....... 

Female genitalia with modified papilla analis (sclerotized and of variable shape, 
rounded, folded or digitate) (Text-figs 17, 18, 19) 

Female genitalia with unmodified papilla analis (membranous and of rounded 
shape) (Text-figs 15, 16) . M. galatheae bechuana subsp. n. (part) 

Fore wing upper surface with costal region pink ...... 

Fore wing upper surface with costal region white, either immaculate or with 
costal margin pink .... M. cruentata (Moore) (part) 

Female genitalia without terminal spines on the papilla analis (Text-fig. 52). 
(From north India, Nepal, Tibet or China) M. cruentata (Moore) (part) 

Female genitalia with terminal spines on the papilla analis (Text-figs 46 to 49). 
(From India or Africa) . . M. galatheae galatheae (Wallengren) 

Fore wing with areole .......... 

Fore wing without areole .......... 

Madagascar, African, Arabian or Iranian species. . . . . . 

Indian species ............ 

Madagascan species. 

Fore wing upper surface light yellow with a wide pink longitudinal band 
occupying nearly the whole costal region (PI. 2, fig. 132). 

M. prochaskai (Viette) (part) 

African, Arabian or Iranian species ........ 

Fore wing upper surface with a light to dark brown lower central longitudinal 
streak, sometimes distad-splayed (PI. 9, fig. 218 and PI. 10, fig. 226) . 

Fore wing upper surface without a light to dark brown lower central longitudinal 
streak ............. 

Fore wing upper surface with a brown discocellular spot as in PI. 9, fig. 218. 

M. bimaculata nigrijasciuta (Hampson) (part) 

Fore wing upper surface without a discocellular spot ..... 

Fore wing upper surface with brown postmedial and terminal dots (PI. 10, 
fig. 230) ..... M. perstriata zernytamsia (Berio) 

Fore wing upper surface with or without brown postmedial dots; brown 
terminal dots absent . . M. perstriata fuscostriata (Brandt) (part) 



(P- 49) 



42 
45 

43 



(p- 


46) 




44 


(p- 


47) 


(p. 


47) 


(p- 


40) 




46 




63 




47 




56 



(P- 


34) 
48 




49 




5i 


(P- 


87) 
50 


(P- 


96) 


(P- 


96) 



orbicular dot 




basal dash 

Figs 25-26. Fore wing markings of Masalia disticta (25) and M. albipuncta (26) 



18 P. R. SEYMOUR 

51 (48) Female genitalia with unmodified papilla analis (membranous and of rounded 

shape) (Text-figs 15, 16) ......... 52 

— Female genitalia with modified papilla analis (sclerotized and of variable shape, 

rounded, folded or digitate) (Text-figs 17, 18, 19). 

Fore wing upper surface from pale to greyish orange, or pastel red through 
brownish orange to dull red. Anal region immaculate or lightly suffused with 
greyish brown (PI. 7, fig. 187) . M. flavistrigata (Hampson) (part) (p. 65) 

52 (51) Female genitalia with the surface of the papilla analis spiculate (striate 

appearance) (Text-fig. 89). 

Fore wing upper surface orange with light brown to brown markings. 

M. nubila (Hampson) (p. 77) 

— Female genitalia with the surface of the papilla analis not spiculate . . 53 

53 (52) Hind wing upper surface light brown to brown. 

Fore wing upper surface light orange irregularly irrorate with brown. 

M. mittoni (Pinhey) (p. 82) 

— Hind wing upper surface yellowish white ....... 54 

54 (53) Fore wing upper surface with costal and anal regions light yellow or dull red to 

greyish pink; central region pale yellow, irrorate with dark brown between 
veins, as in PI. 3, figs 148, 149 M. galatheae bechuana subsp. n. (part) (p. 46) 

— Fore wing upper surface not of this colour and pattern . . . . 55 

55 (54) Fore wing upper surface with an oblique to longitudinal reddish brown to 

brown dash near the base of the anal region (Text-fig. 26). 

M. albipuncta (Hampson) (p. 73) 

— Fore wing upper surface without an oblique to longitudinal reddish brown to 

brown dash near the base of the anal region M. quilengesi sp. n. (part) (p. 74) 

56 (46) Female genitalia with modified papilla analis (sclerotized and of variable shape, 

rounded, folded or digitate) (Text-figs 17, 18, 19) . . . . -57 

— Female genitalia with unmodified papilla analis (membranous and of rounded 

shape) (Text-figs 15, 16) ......... 58 

57 (56) Fore wing upper surface orange or greyish orange marked with brownish 

orange. Female genitalia with the dorso-lateral surface of the papilla analis 

not sericate (Text-fig. 91) ..... M. dora Swinhoe (p. 78) 

— Fore wing upper surface yellowish white to pale orange, often finely irrorate with 

brown (a brown discocellular spot and brown central longitudinal streak may 
be present). Female genitalia with the dorso-lateral surface of the papilla 
analis sericate (Text-fig. 104) . . M. bimaculata bimaculata (Moore) (p. 87) 

58 (56) Female genitalia with papilla analis of right-angle triangular shape (Text- 

fig- 93)- 

Fore wing upper surface either light greyish or brownish orange. 

M. tosta Moore (part) (p. 80) 

— Female genitalia with papilla analis not of right-angle triangular shape . . 59 

59 (58) Fore wing upper surface yellowish white to pale yellow with a series of brown 

postmedial and terminal dots and a poorly differentiated brown lower central 
longitudinal streak . . M. perstriata perstriata (Hampson) (p. 94) 

— Fore wing upper surface not marked in this way ...... 60 

60 (59) Fore wing upper surface with the costal and anal regions light orange to 

pastel-red; central region yellowish white to pale yellow, streaked with light 
orange or pastel-red (a light brown to brown lower central longitudinal 
streak may also be present) . . M. beatrix beatrix (Moore) (part) (p. 59) 

— Fore wing upper surface with the costal, central and anal regions either pale 

light yellow or greyish orange ........ 61 

61 (60) Hind wing upper surface yellowish white with proximad light brown suffusion. 

M. semifusca sp. n. (p. 79) 

— Hind wing upper surface concolorous ........ 62 



REVISION OF GENUS MASALIA 19 

62 (61) Male genitalia with 40 or more closely packed spicules on the scobinate bar. 
(Text-fig. 99) ...... M. modesta (Moore) (p. 84) 

Male genitalia with 20 or fewer sparsely scattered spicules on the scobinate bar 

(Text-fig. 95) ..... M. artaxoides (Moore) (part) (p. 81) 

63 (45) African species ........... 64 

- Indian species ............ 69 

64 (63) Fore wing upper surface pastel yellow with a greyish pink to dull red oblique 
dash extending from the apex to the central region (PI. 2, figs 133, 134). 

M. sublimis (Berio) (p. 35) 
Fore wing upper surface not of this colour and pattern .... 65 

65 (64) Fore wing upper surface with a brown to dark brown lower central longitudinal 
streak (Text-fig. 3) .......... 66 

- Fore wing upper surface without a brown to dark brown lower central 
longitudinal streak .......... 67 

66 (65) Fore wing upper surface with the costal and anal regions pale to dull red; 
central region yellowish white to light yellow with the red colouring occa- 
sionally extending in from the costal and/or anal region; with or without 
brown irroration . . . M. bimaculata nigrifasciata (Hampson) (p. 87) 

- Fore wing upper surface with the costal, central and anal regions white, pale 
yellow, greyish orange or reddish grey, irrorate with brown. 

M. bimaculata cornia subsp. n. (part) (p. 90) 

67 (65) Female genitalia with terminal spines on the papilla analis (Text-fig. 19). 
Fore wing upper surface pale to greyish orange with a faint brown disco- 
cellular spot . ..... M. flavocarnea (Hampson) (p. 92) 

Female genitalia without terminal spines on the papilla analis ... 68 

68 (67) Male genitalia with a cornutus. Fore wing upper surface yellowish white to 
pale or greyish orange . . M. bimaculata cornia subsp. n. (part) (p. 90) 

- Male genitalia without a cornutus. Fore wing upper surface greyish orange, 
greyish red, pale red, or reddish grey. 

M. bimaculata pluritelifora (Berio) (part) (p. 90) 

69 (63) Fore wing upper surface with brown postmedial streaks on veins M\ to Cu\ a , 

as in PI, 7, fig. 195 ...... M. uncta (Swinhoe) (p. 70) 

- Fore wing upper surface without brown postmedial streaks on veins M\ to Cwi a . 70 

70 (69) Female genitalia with modified papilla analis (sclerotized and of variable shape, 

rounded, folded or digitate) (Text-figs 17, 18, 19). 

Fore wing upper surface brownish orange; anal region irrorate with brown, 
otherwise immaculate ..... M. metaphaea (Hampson) (p. 82) 

- Female genitalia with unmodified papilla analis (membranous and of rounded 

shape) (Text-figs 15, 16) ......... 71 

71 (70) Fore wing upper surface bicolorous, light yellow with a brownish outer margin. 

M. albicilia (Hampson) (p. 72) 

- Fore wing upper surface concolorous, light yellow or greyish orange. 

M. artaxoides (Moore) (part) (p. 81) 

DESCRIPTIONS OF THE SPECIES AND SUBSPECIES 

Masalia philbyi (Brandt) comb. n. 
(Text-figs 27-29; PI. 1, figs 114-117; Map 1) 

Timora philbyi Brandt, 1941 : 853. LECTOTYPE <J, Iran (NR, Stockholm), here designated 

[examined.]. 
Timora philbyi nuristana Boursin, i960 : 151. Holotype $, Afghanistan (ZSBS, Munich) 

[examined]. Syn. n. 



P. R. SEYMOUR 



Timora philbyi arabica Boursin, i960 
[examined]. Syn. n. 



152. Holotype $, Saudi Arabia (ZSBS, Munich) 



Antenna with flagellar segments sexually dimorphic (Text-figs 27, 28). Proboscis quotient 
4 to 5. Fore wing with areole; length, <J (21), 13-9-17-6 (lectotype 16-6), $ (18), 14-4-18-3. 
Wing-pattern as in PI. 1, figs 114-117. Fore wing upper surface with ground colour light 
yellow to brownish orange; longitudinal streaks white, either faint or distinctly outlined with 
brownish orange or light brown. Hind wing upper surface white, immaculate or irrorate with 
brownish orange. 

Genitalia, q* scobinate bar and cornutus as in M.fissifascia (Text-fig. 31). $ papilla analis 
finely spiculate, but simple (Text-fig. 29). 

Material examined. 

Timora philbyi, LECTOTYPE, here designated, Iran : Laristan, Strasse Bender- 
Abbas-Sardabad Sardze, about 200 m, <$, xi(mid).i937 {Brandt), in NR, Stockholm; 
paralectotype, Iran: Baloutchistan, Strasse Tchahbahar-Iranchar, Tahte-Malek, 
750 m, <J, iv(early).i938 {Brandt), in NR, Stockholm. Timora philbyi nuristana, 
holotype, Afghanistan: Asmar, Kunartal, 900 m, $, 3.^.1953, in ZSBS, Munich. 
Timora philbyi arabica, holotype, Saudi Arabia: El Riad, $, 6.iii.i958 {E. Diehl), 
in ZSBS, Munich; paratype, Saudi Arabia: El Riad, $, 11. hi. 1958 {E. Diehl), in 
ZSBS, Munich. 

Saudi Arabia: Buraiman, 1 $ {Wiltshire); Dawadami, 1 g, 2 $, 8.iii.i935; Jidda, 
I <j>, 13.ii.1929; 2 $, 28-31.1.1930; 1 $, 1 $, 26.xii.1934; Khafs, 1 $, 26.ii.1935; Marrat, 
3 <?, 6.UI.1935; Mecca, 1 <J, 7.xii.i934; 1 $, 15.il. 1934; Shaib Sudair, 1 <J, 6.11.1935; 
Sir, Khufuiflya, 2 <$, 7.iii.i935; Usba, 3 <$, 1 $, 5.iii.i935 (all coll. H. St. J. B.Philby); 
E. Ryadh, 1 $, 13.iii.1958 {E. Diehl), in ZSBS, Munich; Afghanistan: Sarobi, 
1100 m, 9 (J, 2 $, 15.iv.-10.iv.1961 {G. Ebert), in ZSBS, Munich. 



dorsal 





Figs 27-28. M. philbyi, antennal segments, lateral view. 27, $. 28, <j>. 



REVISION OF GENUS MASALIA 21 

Distribution (Map i). Arabia, southern Iran and east Afghanistan. 

Remarks. The species is readily distinguished from M. fissifascia, the only 
other species in the group, by the difference in fore wing colour, light yellow to 
brownish orange in M. philbyi, greyish red in M. fissifascia. 

The type-specimens of philbyi, nuristana and arabica, differing slightly from one 
another in pattern and colour, lie within a range of continuous variation likely to be 
found in any one locality. 

Masalia fissifascia (Hampson) comb. n. 
(Text-figs 30, 31; PI. 1, figs 118, 119; Map 1) 
Titnora fissifascia Hampson, 1903 : no. 

Antenna with flagellar segments sexually dimorphic, as in philbyi (Text-figs 27, 28). Pro- 
boscis atypically long, quotient 4 to 5. Fore wing with areole present. 

Genitalia. $ scobinate bar and cornutus as in Text-fig. 31; $ papilla analis finely spiculate, 
but simple (Text-fig. 30). 

M . fissifascia can be separated from M . philbyi, the only other species within the 
group and the only one with which it is likely to be confused, on the difference in 
fore wing ground colour. In M. fissifascia the ground colour is greyish red, in 




Fig. 29. M. philbyi, $, papilla analis. 



22 P. R. SEYMOUR 

M. philbyi light yellow to brownish orange. M. fissifascia is also smaller in size, 
though overall ranges of the species overlap. Two subspecies are recognized, 
M. f. fissifascia from Ethiopia, Somali Republic and Kenya, and M. f. roseata from 
Sudan. Subspecies fissifascia can be separated from roseata on the light brown 
outlining of the forked, lower central longitudinal streak, absent in roseata. 




(o) philbyi 

^^ fissifascia fissifascia 

A fissifascia roseata 



Map i. Distribution of species and subspecies of the fissifascia-gToup. 



REVISION OF GENUS MASALIA 



23 



Masalia fissifascia fissifascia (Hampson) 

(PL 1, fig. 118; Map 1) 

Timora fissifascia Hampson, 1903 : no. Holotype $, Kenya (BMNH) [examined]. 

Fore wing length $ (5), 12-4-13-3 (holotype 13-3), ? (6), i6-6-i5-i. Wing pattern as in PI. 1, 
fig. 118. Fore wing upper surface with ground colour greyish red; radial streaks, lower central 
longitudinal streak and anal longitudinal streak white, outlined with light brown. The white 
in the anal longitudinal streak is occasionally missing but more often represented by a short 
proximal line. Hind wing upper surface white, immaculate or irrorated with light brown. 

Material examined. 

Holotype, [Kenya] B.E. Africa: [Kibaoni] Kibauni lC £ 4.X11.1898 (R. Crawshay). 

Ethiopia: Dire Daoua, 1 $, ix.1935 (H. Uhlenhuth); Somali Republic: Buran, 
io°i3'N, 48°47'E, 3000 ft, 1 9, ix-x.1929 (C. L. Collenette) ; Hargeisa, 4300 ft, 1^,1$, 
v.1929 (M. Portal-Hyatt) ; Kenya: Kedai: 1 $, xii.1911 {Feather); Kibwezi, 1 $, 
xii.1921 (Feather); Nairobi, 1 $, iv.1927 (D. H. Hopkins); South Kavirondo, Suna, 
2 c$, 1 $, iii-iv.1932 (W. Feather). 











■■-4lr 





31 



30 



Figs 30-31. M. fissifascia, genitalia. 30,$. 31, $. 



24 P. R. SEYMOUR 

Distribution (Map i). Ethiopia, Somali Republic and Kenya. 

Remarks: The light brown outlining of the forked lower central longitudinal 
streak separates subspecies fissifascia from roseata. 

Masalia fissifascia roseata (Pinhey) comb, et stat. n. 

(PI. i, fig. 119; Map 1) 

Timora roseata Pinhey, 1956 : 13. Holotype <J, Sudan (BMNH) [examined]. 

Forewing length $ (2), 13-2-14-0 (holotype), $ (2), 12-3-13-2. Wing pattern as in PI. 1, 
fig. 119. Fore wing upper surface with ground colour pale to greyish red, longitudinal streaks 
white. Hind wing upper surface white. 

Material examined. 

Holotype, Sudan: Showak, $, ix.1949 (E. Wilson). Paratype, Sudan: Showak, 
$, ix.1949 (E. Wilson). 

Sudan: Ed Darner, Hudeiba, 1 g, 8.viii.i962 (R. Remane), in ZSBS, Munich; 
Blue Nile Province, Wad Medani, 1 £, 2.viii.i962 (R. Remane), in ZSBS, Munich. 

Distribution (Map 1). North-east Sudan. 

Remarks. The absence from M. f. roseata of brown outlining to the longitudinal 
streaks distinguishes it from the nominate subspecies. 



Masalia decorata (Moore) comb. n. 

(Text-figs 32-36; PI. 1, figs 120-125; Map 2) 

Pradatta decorata Moore, 1881 : 365. 

M. decorata is closely allied to M. leucosticta. The absence from M. decorata of 
pink in the proximal to mid costal region of the fore wing distinguishes it from 
M. leucosticta. 

Three subspecies are recognized, M. d. decorata from Afghanistan, India and 
Ceylon, M. d. metarhoda from Africa and M. d. albiseriata from Africa and Saudi 
Arabia. The subspecies are separable on the following genitalic differences in the 
male. 

cornutus of aedeagus scobinate bar of aedeagus 
M. d. decorata present present 

M. d. albiseriata absent present 

M. d. metarhoda present absent, or if present poorly 

differentiated 

In addition the two African subspecies can be separated on differences in fore 
wing pattern and hind wing colour. The fore wing postmedial white dots or dashes 
in M. d. albiseriata lie along the proximal margin of the postmedial band (Text-fig. 33) 
whilst in M. d. metarhoda they lie more-or-less centrally within it (Text-fig. 32). 
In M. d. albiseriata the hind wing upper surface is white to light yellow, in M. d. 



REVISION OF GENUS MASALIA 



25 



metarhoda greyish red, or greyish to brownish orange. Position of the postmedial 
white dots in M. d. decorata is variable, ranging from the condition found in M. d. 
albiseriata to that in M. d. metarhoda. 

The possibility that M. d. albiseriata and M. d. metarhoda are biologically isolated 
is raised by the distribution of M . d. albiseriata and M. d. metarhoda in the Senegal- 
Gambia region, particularly the record of M. d. albiseriata and M. d. metarhoda from 
Dakar, caught on the same day, for no intermediate material was included. Their 
occurrence together cannot be satisfactorily explained, and more material and 
breeding experiments are needed to resolve the problem. However if M. d. albiseriata 
and M . d. metarhoda are biologically isolated they may well be linked as subspecies 
through M. d. decorata. The closer affinities of M . d. albiseriata and M. d. metarhoda 
to M. d. decorata, than to each other and their geographical isolation from M. d. 
decorata suggests an instance of a 'ring of races'. 

Masalia decorata decorata (Moore) 

(Text-fig. 34; PI. 1, figs 120-122; Map 2) 

Pradatta decorata Moore, 1881 : 365. Lectotype $, India (BMNH) [examined]. 
Timora decorata (Moore) Hampson, 1903 : 106. [Lectotype designated.] 

Fore wing with areole; length <$ (57), 9-0-13-6 (lectotype io-i), $ (22), 10-0-12-8. Wing 
pattern as in PI. 1, figs 120-122. Fore wing upper surface with ground colour light yellow; 
transverse and longitudinal markings pastel-red to pink, antemedial dots, when present, and 
postmedial dots white. Hind wing upper surface white to pale yellow. 

Genitalia. <J scobinate bar and cornutus as in Text-fig. 34. $ papilla analis simple. 

Material examined. 

Lectotype, designated by Hampson, [India: Uttar Pradesh,] Saidabad, <$; 
paralectotypes, [Uttar Pradesh], Saidabad, 1 <$ (Atkinson); [U.P.], Allahabad, 1 £ 
[(Hellard); [U.P.] N.W. Provinces, [Mainpuri] Manpuri, 2 <$, 1 $; Deccan, 1 $ {Day); 
[?]; Sind Valley, 1 <J (Atkinson), in MHNU, Berlin. 

Afghanistan: Bashgultal, 1300 m, 1 $, 15.vii.1952 (/. Klapperich); Sarobi, 
1100 m, 1 $, 28.vi.1956 (H. G. Amsel), both in ZSBS, Munich. India: Gujarat, 




Figs 32-33. 32, M. decorata metarhoda, fore wing. 33, M. albiseriata fore wing. 



26 



P. R. SEYMOUR 



Deesa, i <J, i $, viii.1890; 2 ^, 1 ?, x.1899; Himachal Pradesh, Dharmsala, 1 6*; 
Jammu & Kashmir, Srinagar, 5200 ft, 1 $, vii.1892; 1 $, 3.viii.i892; 2 <$, 6.viii.i892; 

1 6*, 10.viii.1892; 5 cJ, 1 $, n.viii. 1892; 3 (J, 1$, 12.viii.1892; 1 (J, 1 $, 14.viii.1892; 

2 (J, 1 $, x.1899; Maharashtra, Nagpur, 5 <$; Mysore, Belgaum, 1 $ (T. R. Bell); 
Uttar Pradesh, Meerut, 1 $, 20.vii.1906 (C. #. Ward) ; West Bengal, Darjeeling, 1 <$, 
{Lidderdale) ; W.B., Calcutta, 1 q*> 1 $; ?, Umballa, 1 <$, 21. ix. 1903, (C. i/. Ward). 
Sikkim: 1 $. Ceylon: Kaits, 1 <$, iii.1901; Puttalam, 7 £, 3 $; 6 o*> 1 $. Burma: 

Distribution (Map 2). Afghanistan, India, Ceylon and Burma. 

Remarks. M. ^. decorata can be separated from M. d. albiseriata and M. d. 
metarhoda on difference in male genitalia. In the nominate subspecies the cornutus 
and scobinate bar are both present, in M. d. albiseriata and M. d. metarhoda one or 
other of these structures is absent; in albiseriata the cornutus, in metarhoda the 
scobinate bar. 




^ decorata decorata 
^ decorata albiseriata 
I decorata metarhoda 



Map 2. Distribution of subspecies of M. decorata. 



REVISION OF GENUS MASALIA 27 

Masalia decorata albiseriata (Druce) comb, et stat. n. 
(Text-figs 33, 35; PL i, fig. 123; Map 2) 

Timora albiseriata Druce, 1903 : 200. LECTOTYPE o". Gambia (BMNH), here designated 

[examined]. 
Timora buchanani Rothschild, 1921 : 160. Holotype $, Niger (BMNH) [examined]. Syn. n. 

Fore wing with areole; length <$ (9), 8-9-10-7 (lectotype 8-9), $ (10), 9-2-11 -2. Wing pattern 
as in PI. 1, fig. 123. Fore wing upper surface with ground colour light yellow; transverse and 
longitudinal markings pastel-red to pink; antemedial dots when present, and postmedial dots, 
white. Hind wing upper surface white to light yellow. 

Genitalia. $ atypical in that the cornutus is absent; scobinate bar and proximal part of 
vesica as in Text-fig. 35; ? with papilla analis simple, as in M. d. decorata. 

Material examined. 

Timora albiseriata, LECTOTYPE, here designated, Gambia, $ (A. Moloney). 
Paralectotype, Gambia, 9 (A. Moloney). Timora buchanani, holotype [Niger:] 
Azzal, North of Agades, $, 14.vii.1920 (A. Buchanan). Paratypes. [Niger:] 
Azzal, 1 $, 13.vii.1920; 1 <3\ 14.vii.1920; 2 J, 1 $, 15.vii.1920. 

Senegal: Dakar, 1 <J, 12.ix.1956 (C. Rungs), in MNHN, Paris; Kaolack, 2 $ 
{LeMoult). Kenya: Wajir, 1 $, iv.1958 {Hutchinson). [?] : Saloum, 1 <J, v.1926. 
Saudi Arabia: Jidda, 1 $, 3.U.1930; 1 ?, 8.1V.1930; itf, 1 $, 26.V.1930 (all//. St J. B. 
Philby) ; Mujaririma, 1 $, 1 $, L1945 (B. P. Uvarov). 

Distribution (Map 2). Senegal, Gambia, Ghana, Niger, Sudan and Saudi 
Arabia. 

Remarks. The absence of a cornutus in the male of M. d. albiseriata readily 
separates it from M. d. metarhoda and M. d. decorata; in each of the latter two a 
cornutus is present and well developed. M. d. albiseriata can also be distinguished 
from M. d. metarhoda on difference in fore wing pattern and hind wing colour, the 
postmedial white dots of the fore wing in M. d. albiseriata lie along the proximal 
margin of the postmedial band, in M. d. metarhoda they lie more-or-less centrally 
within it. The hind wing colour of M. d. albiseriata is white to light yellow, that of 
M. d. metarhoda greyish red, or greyish to brownish orange. 



Masalia decorata metarhoda (Druce) comb, et stat. n. 

(Text-figs 32, 36; PL 1, figs 124, 125; Map 2) 

Timora metarhoda Druce, 1903 : 201. Lectotype $, Gambia (BMNH) [examined]. 
Timora metarhoda Druce; Hampson, 1903 : 106. [Lectotype designated.] 

Fore wing with areole; length 5* (20), 9-2-11-2, $ (27), 9-8-12-2 (lectotype). Wing pattern 
as in PL 1, figs 124, 125. Fore wing with upper surface ground colour light to greyish yellow; 
transverse and longitudinal markings pastel-red to pink; antemedial dots, when present, and 
postmedial dots, white. Hind wing with upper surface greyish red or greyish to brownish 
orange. 



28 



P. R. SEYMOUR 




Figs 34~3 6 - M. decorata subspecies, $, scobinate bar and cornutus. 34, M. d. decorata. 
35, M . d. albiseriata. 36, M. d. metarhoda. 



REVISION OF GENUS MASALIA 29 

Genitalia. <$ with scobinate bar absent or poorly differentiated. Distal end of aedeagus 
and proximal part of vesica as in Text-fig. 36. $ with papilla analis simple, as in M . d. decorata. 

Material examined 

Lectotype, designated by Hampson, Gambia, $ (A. Moloney). 

Senegal: Dakar, 3 <$, 1 $, 12.ix.1956 (C. Rungs), in MNHN, Paris; Kaolack, 1 <$ 
(LeMoult) ; 3 $ (G. Melon) ; N'dande, 1 <J, 26.viii.1951 (B. Boniface), in MNHN, Paris. 
Ghana: Northern Territories, Navaro, 1 $; 1 $, x.1923 (both A. W. Cardinall). 
? Niger: Baban Tubki, south of Zinder, 1 $, 13. ix. 1920 (A. Buchanan). Niger: 
Damergou, Bande, 4 $, 7 $, 16. ix. 1920; Kaleloua, 2 <£, 3 $, 8.ix.i92o; Makochia, 
6 (J, 8 $, 15. ix. 1920; Tanout, irf, 6.ix.i92o; Zinder, 1 $, 11. ix. 1920 (all ^4. Buchanan). 
Sudan: Khartoum, 1 <J, 5_ix.i928 (//. W. Bedford); White Nile, 12-15 N., 1 $ 
(Yardley). 

Distribution (Map 2). Senegal, Gambia, Ghana, Niger and Sudan. 

Remarks. The absence of, or a poorly differentiated, scobinate bar in the male 
of M. d. metarhoda distinguishes it from both M . d. albiseriata and the nominate 
subspecies. In these latter two the scobinate bar is well developed. M. d. meta- 
rhoda can also be distinguished from M. d. albiseriata on difference in fore wing 
pattern and hind wing colour; the postmedial white dots of the fore wing in M. d. 
metarhoda lie more-or-less centrally within the postmedial band (Text-fig. 32), in 
M. d. albiseriata they lie along the bands proximal margin (Text-fig. 33). Hind 
wing colour of M . d. metarhoda is greyish red, or greyish to brownish orange, that of 
M. d. albiseriata, white to light yellow. 



Masalia leucosticta (Hampson) comb. n. 
(Text-figs 37, 38; PI. 2, figs 126-130; Map 3) 

Timora leucosticta Hampson, 1902 : 256. 

M. leucosticta can be distinguished from the closely allied M. decorata by the 
difference in colour of the proximal area of the costal region, pink in M. leucosticta, 
pale to olive yellow in M. decorata ; and from M. funebris, to which M. leucosticta 
joiceyi shows particularly close affinities, on difference in hind wing colour. 

Three subspecies are recognized: leucosticta, vinula and joiceyi. The three are 
separable on fore wing differences and distribution. M. 1. vinula and M. I. joiceyi 
have contiguous ante- and postmedial markings distinguishing them from M. I. 
leucosticta in which the markings are not contiguous. The distribution of M. I. 
vinula is northern Tanzania and northward to Ethiopia ; that of M . I. joiceyi, southern 
Tanzania and westward to the southern Congo (Kinshasa). The areole, usually 
present in M. 1. vinula, is rarely present in M. I. joiceyi. 






30 



P. R. SEYMOUR 



Masalia leucosticta leucosticta (Hampson) 
(Text-fig. 38; PI. 2, figs 126-128; Map 3) 

Timora leucosticta Hampson, 1902 : 256. Lectotype o\ Botswana (BMNH) [examined]. 

Timora leucosticta Hampson; Hampson, 1903 : 107. [Lectotype designated] . 

Timora continuata Grunberg, 1910 : 126. LECTOTYPE $, South West Africa (MNHU, 

Berlin), here designated [examined]. Syn. n. [Gaede 1935 : 105 refers to continuata as a 

form, both of metarhoda and leucosticta]. 

Fore wing with areole usually present; length £ (66), 10-9-13-4 (lectotype 12-5), $ (51), 119- 
14 o. Wing pattern as in PI. 2, figs 126-128. Fore wing upper surface with ground colour 
light yellow, transverse and longitudinal markings pink to greyish pink, postmedial dots and 
when present, antemedial dots white. Hind wing with upper surface white to pale yellow, 
immaculate or with subterminal or whole area behind subterminal region irrorate with greyish 
brown. 

Genitalia, q? scobinate bar and cornutus (Text-fig. 38). 9 with papilla analis simple. 






Material examined. 

Timora leucosticta, lectotype, designated 
Country, $ (F. D. Lugard). 



by Hampson, [Botswana] N 'Garni 
Timora continuata, LECTOTYPE, here designated, 




,\7 




37 




Figs 37—38. M. leucosticta subspecies, q\ scobinate bar and cornutus. 37, M. l.joiceyi. 

38, M. I. leucosticta. 



REVISION OF GENUS MASALIA 



3i 



South West Africa: Okahandja, [Windhoek] 'Windhuk', g, 24.ii.1909 (S. G. 
Seewald), MNHU, Berlin. South West Africa: Hoffnung (east of Windhoek), 
1850 m, 5cJ, 9. i. 1934; 40^, 38 $, 10. i. 1934; 1 <$, 30.1.1934 (all K.Jordan) ; Okahandja, 
2 <$, 20-26.1.1928; 1 J, 27.i-2.ii.1928; 1 (J, 1 $, 3-16. ii. 1928; 2 (J, 2-18. iii. 1928; 2 J, 
1 $, 19-29. iii. 1928 (all i?. £. Turner); 1 <J, ii.1935 (F. Gaerdes); 1 (J, 27. ii. 1945; 1 (J, 
24.iii.1947; 1 o\ 24.V.1947; 2 $, 25. iii. 1954; 1 $, 5.iv.i953 (all F. Gaerdes), in ZSBS, 
Munich; Windhoek, 1 $ (Knier), in ZSBS, Munich; 1650 m, 1 J, 2 $, 16. i. 1934; 2 $, 
19. i. 1934 (all A'. Jordan); Tanzania: Kongwa, 2 <J, 23.iv.1917 (A. Loveridge); 
District of Great Craters, 1 <$, ii-iii.1921 (T. A. Bams) ; Mamboia, 1 o (Baxter). The 
following specimens are intermediate between M. 1. leucosticta and M. I. vinula, but 
placed with M. 1. leucosticta: Kondoa Irangi (dry sandy country), 1 $, iii.1921 (T. A. 
Barns) ; Shinyanga, 1 $, i.1957 [Croft) ; Mwanza, 1 $, xii.1925-i.1926; Arusha District, 
Odeani Crater (dry thorn bush country), 1 $ (T. A. Barns). 

Distribution (Map 3). South West Africa, Botswana and Tanzania. 




^P leucosticta leucosticta 

{*J leucosticta leucosticta (intermediate) 

A leucosticta vinula 

A leucosticta vinula (intermediate) 

\< leucosticta joiceyi 



^ 



Map 3. Distribution of subspecies of M. leucosticta. 



32 P. R. SEYMOUR 

Remarks. The non-contiguous ante- and postmedial pink markings on the fore 
wing of M. 1. leucosticta separate it from M. I. vinula and M. I. joiceyi ; in these latter 
two subspecies the markings are contiguous. 



Masalia leucosticta vinula (Berio) comb, et stat. n. 
(PI. 2, fig. 129; Map 3) 

Timora vinula Berio, 1943 : 182. Holotype <J, Ethiopia: Elaberet, 17. ix. 1938 (G. Vaccaro) 
(Berio Coll., MCSN, Genoa). 

Fore wing with areole usually present; length <J (42), 10-4-13-8, $ (13), 10-5-14-2. Wing 
pattern as in PI. 2, fig. 129. Fore wing upper surface with ground colour light yellow; post- 
medial dots, and when present antemedial dots, white; other markings pink to greyish pink. 
Hind wing with upper surface pale yellow, immaculate, or pale yellow with subterminal or 
whole area behind subterminal region irrorate with greyish brown. 

Genitalia. <J with scobinate bar and cornutus as in M. 1. leucosticta (Text-fig. 38) ; $ papilla 
analis simple, as in M. I. leucosticta. 

Material examined. 

[Tanzania:] Miriu River, 1 $, iii.1925 (D. Hopkins); Musoma, Banagi Hill, 2 <$, 
1 $, iii.1957. Kenya: Isiolo, 1 <$, iv.-v.1951; 2 <$, iv.1954 (/. Adamson); Makindu 
(south of Nairobi), 1 $, iii.1927 (W. Feather); Masongoleni, 1 $, 25.iii.1911; 1 <$, 
30.iv.1911; 1 $, 25.ix.19n; 1 g, 30.iv.19n (all W. Feather); Mombasa, 2 <$, vi.1916 
(van Someren); Kibwezi, 2 $, 9.xii.i9i6; 8 £, 12.xii.1916; 1 <$, 21.xii.1916; 1 $, 
23.iv.1917; 1 <J, 11.xii.1918; 6 <$, 1 $, 19.xii.1918; 5 <$, 22.xii.1918; 2 (J, 1 $, 23.xii. 
1918; 2 $, 13.iv.1919; 1 <$, 2.V.1919; 1 <$, 2.V.1919; 1 <$, 2.V.1919; 1 g, 4.V.1919; 1 $, 
xii.1920; 1 (J, 3 $, iv.1922 (all W. Feather). The following specimens show variation 
in pattern between M. 1. leucosticta and M. 1. vinula but are placed with M. I. vinula. 
Ethiopia: Harar, 1 $, 24.iv.1939 (R. E. Ellison); Somali Republic: Hargeisa, 
4300 ft, 1 (J, 1 $, v.1929 (M. Portal-Hyatt). 

Distribution (Map 3). Tanzania, Kenya, Ethiopia and Somali Republic. 

Remarks. The contiguous ante- and postmedial pink markings on the fore wing 
of M. 1. vinula distinguish it from M. 1. leucosticta. Differences between M. I. vinula 
and M. I. joiceyi are less marked ; joiceyi tends to be of larger size and more robust 
in appearance and the areole, in contrast with vinula, is rarely present. M. I. vinula 
and M. I. joiceyi are separable on distribution, vinula occurring in northern Tanzania 
and northward to Ethiopia, joiceyi occurring in southern Tanzania and westward to 
the southern Congo (Kinshasa). 

Type-specimens have not been examined; determination is based on a paratype 
photograph (Eritrea: Elaberet, <$, i.ix.1938) presented by Dr E. Berio and a speci- 
men from the BMNH Collection identified by him. 



REVISION OF GENUS MASALIA 33 

Masalia leucosticta joiceyi (Prout) comb, et stat. n. 

(Text-fig. 37; PI. 2, fig. 130; Map 3) 
Timor a joiceyi Prout, 1921 : 119. Holotype $, Congo (Kinshasa) (BMNH) [examined]. 
The subspecies is known only from male specimens. 

Fore wing without areole; length $ (4), 13-6 (holotype)-i6-5. Wing pattern as in PI. 2, 
fig. 130. Fore wing upper surface with ground colour yellowish white to pale yellow, markings 
pink to greyish pink; postmedial and antemedial dots white. Hind wing upper surface white, 
with either subterminal or whole area finely to moderately irrorate with greyish brown. 

Genitalia. rf scobinate bar and cornutus as in Text-fig. 37. 

Material examined. 

Holotype, Congo (Kinshasa) : Lufira River near Likasi Copper Mine, 4000 ft, 
<£, 6.xii.i9i8 (T. A. Barns). 

Congo (Kinshasa): Elisabethville, i <$, 16.xii.1954 (C. Seydel). Tanzania: 
Songea, 1 $, 19.1.1933 (R. F. Johnstone) ; ? ; Ningpo, 1 <$, in ZSBS, Munich. 

Distribution (Map 3). Congo (Kinshasa) and Tanzania. 

Remarks. M. 1. joiceyi and M. funebris are closely allied; M. funebris is much 
darker in colour with extensive red areas in the fore wing and with its brown hind 
wings readily distinguished from M. I. joiceyi. The contiguous ante- and post- 
medial pink markings on the fore wing of M. 1. joiceyi distinguish it from M. I. 
leucosticta. M. 1. joiceyi and M. I. vinula are separable on distribution, joiceyi 
occurring in southern Congo (Kinshasa) and southern Tanzania, vinula occurring 
in northern Tanzania and northward to Ethiopia. 

Masalia funebris (Berio) comb. n. 
(Text-fig. 39; PI. 2, fig. 131) 

Timora funebris Berio, 1962 : 125. Holotype <J, Congo (Kinshasa) (MRAC, Tervuren) 
[examined]. 

The species is known only from male specimens. 

Fore wing without areole; length £ (1), 147 (holotype). Wing pattern as in PI. 2, fig. 131. 
Fore wing upper surface with costal and central regions bluish red, finely irrorate with brown; 
basal streak and outer marginal area pale to light orange; anal region brown, post- and ante- 
medial dashes white. Regions not clearly defined. Hind wing upper surface brown. 

Genitalia. <J scobinate bar and cornutus as in Text-fig. 39. 

Material examined. 

Holotype, [Congo (Kinshasa):] [Elizabethville] Elisabethville, $, x.1933 
(C Seydel), in MRAC, Tervuren. 

Distribution. Southern Congo (Kinshasa). 

Remarks. M . funebris is most closely allied to M. sublimis and M. leucosticta 
particularly to M. I. joiceyi ; the brown fore wing markings and brown hind wings of 
funebris, however, readily distinguish it from them. 



34 



P. R. SEYMOUR 



Masalia prochaskai (Viette) comb. n. 
(Text-fig. 40; PI. 2, fig. 132) 

Timora prochaskai Viette, 1957 : 271. Holotype <J, Madagascar: Betioky-Sud, i.1955 
(Prochaska) (MNHN, Paris). 

The species is known only from male specimens. 

Fore wing with areole; length 5* (1), 8-9. Wing pattern as in PI. 2, fig. 132. Fore wing 
upper surface with ground colour light yellow; costal region, orbicular and discocellular spots, 
and post- and antemedial lines, greyish to dull red; post- and antemedial dots white, poorly 
differentiated. Hind wing upper surface pale yellow, suffused with dull red. 

Genitalia. <J scobinate bar and cornutus as in Text-fig. 40. 

Material examined. 

Madagascar: Betioky-Sud, <$, i.1955 (Prochaska), in MNHN, Paris. 

Distribution : Madagascar. 

Remarks. Allied to M. decorata and M. leucosticta. The absence of pink from 
the proximal area of the costal region in M. decorata distinguishes it from 
M. prochaskai. Differences between leucosticta and prochaskai are less marked ; in 
leucosticta the pink longitudinal costal band is comparatively narrow, passing 
through only a part of the costal region, and often broken postmedially. In 




Fig. 39. M. funebris, <J, scobinate bar and cornutus. 



REVISION OF GENUS MASALIA 



35 



prochaskai the band is complete and much wider, occupying nearly the whole costal 
region. 

The single specimen of prochaskai examined was determined by Dr P. Viette. 

Masalia sublimis (Berio) comb. n. 

(Text-fig. 41 ; PI. 2, figs 133, 134) 

Timora sublimis Berio, 1962 : 126. Holotype <$, Congo (Kinshasa) (MRAC, Tervuren) 
[examined]. 

Fore wing without areole; length, <$ (8), 13-8-16-2 (holotype 14-6), ? (1), 15-3. Wing pattern 
as in PI. 2, figs 133, 134. Fore wing upper surface with ground colour pastel-yellow; markings 
dull red to greyish pink. Hind wing upper surface white. 

Genitalia. (J scobinate bar and cornutus as in Text-fig. 41. $ papilla analis simple. 

Material examined. 

Holotype, Congo (Kinshasa) : Kalule Nord, $, i.1934 {Seydel), in MRAC, Tervuren. 

? Angola: 1 cJ, 31.xii.1912 (Rohan-Chabot) , in MNHN, Paris; Angola: i <J, i.1913 
(Rohan-Chabot); Congo (Kinshasa); Elizabethville, 1 <$, 31.xii.1932 (C. Seydel); 
Sandoa, Luiva, 1 <$, 10. iv. 1932 (F. G. Overlaet), both from Mus. Tervuren; Sandoa, 
1 $, ii.1924, Zool. Staatssamml., Munchen; Zambia: Mumbwe, 1 <$, 6.1.1957 
(R. C. Dening); Mwengwa, 1 <J, 18J.1914 (Dollman); Rhodesia: Gwaai River, 
I (J, 17.ii.1921 (C. E. Godman). 




Fig. 40. M. prochaskai, q\ scobinate bar and cornutus. 



36 



P. R. SEYMOUR 



Distribution. Angola, Congo (Kinshasa), Zambia and Rhodesia. 

Remarks. M. sublimis is closely allied to M. leucosticta, M. decorata and 
M. funebris ; the incomplete postmedial band and absence of postmedial white dots 
from the fore wing distinguish it from them. M. sublimis is not likely to be confused 
with any other species. 



Masalia disticta (Hampson) comb. n. 

(Text-figs 25, 42; PI. 2, figs 135-137; M &P 4) 
Timor a disticta Hampson, 1902 : 441. 

M. disticta is separable from other species within the group and genus in possessing 
a white orbicular and white discocellular dot (Text-fig. 25). 

Three subspecies are recognized; the nominate subspecies from South Africa, 
flavirosea from Central Africa and albirosea from East Africa (Map 4). The sub- 
species differ from one another in extent of fore wing pink and yellow colouring. In 
albirosea, known only from the worn type, the costal region is pink, the central and 
anal regions yellowish white. In subspecies disticta, the costal and anal regions are 
pink, the central region pale yellow. In flavirosea all three regions are pink, a pale 
yellow basal dash extending basi-posteriorly to the anal region. 




Fig. 41. M. sublimis, $, scobinate bar and cornutus. 



REVISION OF GENUS MASALIA 37 

Masalia disticta disticta (Hampson) 
(Text-fig. 42; PI. 2, fig. 136; Map 4) 
Timora disticta Hampson, 1902 : 441. Holotype <J, Lesotho (BMNH) [examined]. 

Fore wing with areole present, well developed; length £ (2), 15-3-16-2 (holotype), $ (2), 
16-5. Wing pattern as in PI. 2, fig. 136. Fore wing upper surface with costal and anal regions 
pink, central region pale yellow, orbicular and discocellular dots white. Hind wing upper 
surface white. 

Genitalia, o* scobinate bar and cornutus as in Text-fig. 42. $ papilla analis simple. 

Material examined. 

Holotype, [Lesotho] Basutoland: Masite, <$, 31. 1.1902 (S. Weigall). 
South West Africa: Okahandja, 1 $, 17.1.1958, in ZSBS, Munich; Windhoek, 
1650 m, 1 $, 20. i. 1934; South Africa: Pretoria, 1 <$, 19. i. 1906. 

Distribution (Map 4). South Africa. 

Remarks. The pale yellow central and pink anal region in the fore wing of 
M. d. disticta distinguishes it from M. d. flavirosea (central region pink) and 
M. d. albirosea (anal region white). 




Fig. 42. M. disticta disticta, <$, scobinate bar and cornutus. 



38 



P. R. SEYMOUR 



Masalia disticta albirosea (de Joannis) comb, et stat. n. 

(PI. 2, fig. 135; Map 4) 

Timora albirosea de Joannis, 1913 : 124. LECTOTYPE $, Ethiopia (MNHN, Paris), here 
designated [examined]. 

Fore wing with areole; length q" (i), 147 (lectotype). Wing pattern as in PI. 2, fig. 135. 
Fore wing upper surface with costal region pink, central and anal regions yellowish white, 
orbicular and discocellular dots white. Hind wing upper surface white. 

Genitalia, o" n °t examined but scobinate bar and cornutus almost certainly as in 
M. d. disticta (Text-fig. 42). 

Material examined. 

LECTOTYPE, here designated, [Ethiopia:] Eritrea, $, in MNHN, Paris. 
Distribution (Map 4). East Africa. 




^p disi icta disticta 
^ disticta flavirosea 
I disticta albirosea 



Map 4. Distribution of subspecies of M. disticta. 



REVISION OF GENUS MASALIA 39 

Remarks. The yellowish white central and anal regions in the fore wing of 
M. d. albirosea separate it from the nominate subspecies (anal region pink) and from 
M. d. flavirosea (central and anal regions pink) . 

As a species, M. albirosea was described from two specimens of which only one, 
labelled type, survives. Although in this specimen the pink fore wing markings 
are considerably rubbed, there is no trace of pink from the central or anal regions. 

Masalia disticta flavirosea (Hampson) comb, et stat. n. 

(PI. 2, fig. 137; Map 4) 
Timora flavirosea Hampson, 1903 : III. Holotype o\ Kenya (BMNH) [examined]. 

Fore wing with areole present, well developed; length q* (18), 13-6-1 8 5 (holotype 16-7), 
$ (3), 16-4-17-3. Wing pattern as in PI. 2, fig. 137. Fore wing upper surface with ground 
colour pink, basal dash pale yellow, orbicular and discocellular dots white. Hind wing upper 
surface white. 

Genitalia, o* scobinate bar and cornutus as in M. d. disticta (Text-fig. 42). $ papilla analis 
simple. 

Material examined. 

Holotype, [Kenya:] Machakos, <$, g.xii.i898 (R. Crawshay). 

Angola: Cubal River, 1 <$, iii.1899; Lepi, 3500 ft, 1 <$ (E. Robins); Luimbale, 
1800-1900 m, 7 $, 1 $, 15-20.iii.1934 (K. Jordan) ; Tanzania: Arusha District, 1 $; 
Arusha District, Ngorongoro Crater, 5800-5900 ft, 3^, 1 $, ii-iii.1921 (M. S. Moore) ; 
Musoma, 1 $; Njombe, 6000-6500 ft, 1 $, 25.il. 1952; Kenya: Athi River, 1 <$, 
13.V.1899 (C. S. Bettori); Kikuyu District, 1 <$; Kilindini, 1 J, 27.iv.1899. 

Distribution (Map 4). Central Africa. 

Remarks. M. d. flavirosea can be separated from M. d. disticta and M . d. albirosea 
on the colour difference of the central region of the fore wing : pink in flavirosea, 
yellowish white to pale yellow in disticta and albirosea. 

THE galatheae-cruentata complex 

This complex is made up of several variant lines tentatively arranged into two 
species, M. galatheae and M. cruentata. 

Within the galatheae-group, M. galatheae and M. cruentata are closely allied to 
M. disticta and M. flaviceps. M. galatheae (excepting M. g. bechuana) and 
M . cruentata are separable from M. disticta on differences in the female papilla analis, 
which is modified in M. galatheae and M. cruentata, simple in M . disticta. Separation 
of M. g. bechuana and M. disticta can be made on the absence and presence, respec- 
tively, of a white fore wing orbicular and discocellular dot. With its white fore 
wings M. flaviceps is readily distinguished from M. galatheae and M. cruentata. 

Differences between M. galatheae and M. cruentata are slight. In the fore wing 
pattern of M . cruentata the pink-red costal longitudinal marking is usually narrow 
and the boundaries of this and the pink-red, anal longitudinal marking toward the 
central region, are diffuse. In M. galatheae the pink-red, costal longitudinal marking 



40 P. R. SEYMOUR 

is wide, occupying most or all of the costal region and the boundaries of this and the 
pink-red, anal longitudinal marking, toward the central region, are sharply defined. 
In the female genitalia the papilla analis of M. cruentata is more pointed and it lacks 
the terminal spines found in M. g. galatheae (Text -figs 46-49, 52) ; in the male the 
scobinate bar of M. cruentata has fewer spicules (cruentata 10-40, galatheae 40-150). 

Masalia galatheae (Wallengren) comb. n. 
(Text-figs 43-51; PI. 3, figs 138-149; Map 5) 
Leocyma galatheae Wallengren, 1856 : 58. 

The species is distributed across Africa and central and southern India. 

Two subspecies are recognized, galatheae and bechuana, the latter being restricted 
to a region of southern Africa and structurally separated from the nominate sub- 
species on the difference in the female papilla analis, simple in bechuana, modified 
in galatheae. In the nominate subspecies, variation occurs in size, fore wing shape, 
presence or absence of an areole and in fore wing colour and pattern. Variation 
with respect to these four variables shows a fairly high degree of linkage, though a 
small number of apparently random intermediates do occur. On the basis of linked 
variation a number of forms are recognized; four, each formerly described as species, 
are referred to by their originally proposed names: galatheae, imitata, nigrolineata 
and splendens. 

Masalia galatheae galatheae (Wallengren) 
(Text-figs 43-49; PI- 3. figs 138-144; Map 5) 

Leocyma galatheae Wallengren, 1856 : 58. Holotype $, South Africa (NR, Stockholm) 

[examined] . 
A laria lanceolata Walker, 1865 : 767. Holotype o\ India (BMNH) [examined]. [Synonymized 

by Aurivillius, 1925 : 12.] 
Adisura splendens Druce, 1887 : 685. LECTOTYPE $, Gambia (BMNH), here designated 

[examined]. [Synonymized with lanceolata by Hampson, 1903 : in.] 
Adisura imitata Druce, 1889 : 301. LECTOTYPE <J, 'Costa Rica' [see below] (BMNH), here 

designated [examined]. [Synonymized with lanceolata by Hampson, 1903 : 111.] 
Curubasa depicta Swinhoe, 1891 : 146. Holotype $, India (BMNH) [examined]. [Synony- 
mized with lanceolata by Hampson, 1903 : 111.] 
Timora galatheae (Wallengren) Aurivillius, 1925 : 12. 
Timora nigrolineata Aurivillius, 1925 : 12. Holotype $, Uganda (NR, Stockholm) [examined]. 

Syn. n. 
Timora vittulata Aurivillius, 1925 : 12. LECTOTYPE $, Sudan (NR, Stockholm), here 

designated [examined]. Syn. n. 
Timora lancea Berio, 1953 : 2. Holotype $, Ethiopia: Neghelli, 1441 m, v-vii.1938 (Nicotra) 

(Berio Coll., MCSN, Genoa). Syn. n. 

Fore wing with areole in all forms except splendens; length $ (421), 10-5-18-0 (holotype 13-8), 
$ (198), 11-7-20-0. Wing-pattern as in PL 3, figs 138-144. Fore wing upper surface with 
costal and anal regions pink, red or greyish red, central region white to light yellow; typical 
form and form splendens (PI. 3, figs 138-141) with costal and anal regions pastel pink to greyish 
rose, central region white or yellowish white; form nigrolineata (PL 3, fig. 143) with costal and 



REVISION OF GENUS MASALIA 



4i 



anal regions greyish red to dull red, central region light yellow; form imitata (PI. 3, fig. 142) 
with costal and anal regions red to brownish red, central region pale to light yellow. Hind 
wing upper surface white, immaculate or moderately irrorate with light brown. 

Genitalia, <J scobinate bar and cornutus as in Text-figs 43-45. 9 papilla analis modified; 
terminal spines present (Text-figs 46-49). 




Figs 43-45. M. galatheae galatheae, £, scobinate bar and cornutus. 43 and 45, extremes 
of range found in form galatheae. 44, form imitata. 



42 



P. R. SEYMOUR 







I i 



46 





(■:■■■' 

r 



j ; ' : -^ ; j/ 






* :«/ 



47 



48 




Figs 46-49. M. galatheae galatheae, $, papilla analis. 46 and 47, form imitata. 
48, form depicta. 49, form galatheae. 



revision of genus masalia 43 

Material examined. 

Leocyma galatheae, holotype (labelled Cucullia galathaea Wallengren), [South 
Africa:] Caffraria, <$ (I. Vahlb) ['Kaffern Wahlberg' in original description], in NR, 
Stockholm. Alaria lanceolata, holotype, [India: Madras,] Ccimbatoor, $ 
(M. I. Walhouse). Adisura splendens, LECTOTYPE, here designated, Gambia, 9 
(A. Moloney) ; paralectotype, [Gambia,] i £ (Carter). Adisura imitata, LECTOTYPE, 
here designated, 'Costa Rica' [in fact almost certainly from West or Central Africa, 
the printed label 'Costa Rica Van Pattern' having been mistakenly added to the 
pin], $. Curubasa depicta, holotype, [India: Maharashtra,] Khandala, $ 
(C. Swinhoe). Timor a nigrolineata, holotype, [Uganda]: Sembliki, Lake Edward, 9 
(Swed. Exp. Centr. Africa), in NR, Stockholm. Timora vittidata, LECTOTYPE, 
here designated, Sudan: Renk, Nile, J, in NR, Stockholm; paralectotypes, Sudan: 
['Renk' according to original description] Nile, 2 <J, 1 $ [type-series 3 <$, 1 9> not 
2 (J, I $, as stated in original description], in NR, Stockholm. 

Senegal: Gassane, 1 9. 27.viii.1g07 (W. Riggenbach) ; Kaolack, 1 <$, 4 $, 1909 
(G. Melon); S^dhiou, 2 <$, 2 9. 1917 (H. Castell); Ghana: Aburi, 1 q, i $, 14.x. 1901 
(W. H. Johnstone); Northern Territories, Kete-Krachi, 21 <$, 29 $; Navaro, 1 £, 
viii.1923 (both A. W. Cardinall); Togo: 3 $, 10-14.ix.1893; 1 $, 28. ix. 1893 (both 
L. Conradt); Nigeria: Ogruga [POgrugru], River Niger, 9 <$, 2 $; Agbaja, 1 o*, 
viii-ix.1913 (D. Caton) ; Assaba, R. Niger, 1 $ (Crosse) ; Ilesha, 5 <$ (Humfrey) ; Minna, 
4 <3\ 1-6.X.1910; 3 (J, 1 ?, 8-9.X.1910; 2 <$, 18-19. ix. 1899 (G. Migeod); Cameroun: 
Batouri District, Gadji, 4°3o'N, 4°i5'E, 750 m, 5 <$, 1 $, 1935 (F. G. Merfield) ; 
Gabon: Tchibanga, 1 9> iv.1952 (P. Rougeot); Central African Republic: Fort 
Crampel, 1^,1$; Angola: Ambace, 6 $ (Ansorge); Capelongo-Dongo, 1 J, i.1913; 
Ceramba, 3 <$, 2 $, iii.1903 (W. C. Bell) ; Cubal River, 1 <J, iii.1899 (Penrice) ; Fazenda 
Congula, Amboim District, 7-800 m, 7-22.iv.1934 (K. Jordan); near Lobita, 1 <$ 
(K. H. Cohlan); Luimbale, Mt. Moco, 1800-1900 m, 4 <$, 13.iii.1934, 61 <$, 9 9. 
15-20. iii. 1934, 1 9. 24.iii.1934 (K. Jordan); Lunda, Xa Sengue, 1 J, 2 9. 4-9.iv.1937 
(/I. Exell); Malange, Cocolo, 1 <$, 10.iv.1937 (A. Exell); Quiculungo, 120 km N. of 
Lucala, 800 m, 5 £, 1 9. iv.1936 (R. Braun); Quirimbo, 75 km E. of Amboim, 
300 m, 2 (J, 1 9, 1-6. v. 1934; 15 (J, 26 9, 7-12. v. 1934; 3 <$, 4 9, 13-20. v. 1934 (all 
K. Jordan) ; South Africa : 2 <$ ; Daimana, near Ladysmith, 1 <J (G. A . K. Marshall) ; 
Maritzbourg, 1 9 (Mills); Modderfontein, 5 <$, 1 $, xi.1920 (A. V. Langshaw); 
1 (J, 2 ?, i.1921 (A. V. Langshaw); Natal, 1 <$ (Spitter); 1 £; Duff's Road, 1 <$; 
Pretoria, 2 9; 1 6*> 11-1895; 1 9. 28.x. 1894; 1 $, xi.1894 (all W. L. Distant); Tsomo, 
1 9» (/• H. Bowker); White River, 1 9. xii.1908 (^4. T. Cooke); Weenen, 1 9. 
1 <£, i-ii.1928 (H. P. Thomasset); Zutrsenka, Waterberg District, 1 9. 1. iii. 1899. 
Lesotho: i J, 2 $; Rhodesia: Mashonaland, 2 9 (H. B. Dobbie); 1 $, 16.xii.1897; 
1 9. xii.1900; 4 ^, xii.1904 (all G. A. K. Marshall); Zambia: Chinundo Valley, near 
Lundazi, 1 9, 14.iii.1939 (F. B. Macrae) ; Fort Jameson, 3^ (/. M. Phipps) ; Kaluluma 
Valley, near Lundazi, 1 9, 18.iii.1939 (F. B. Macrae) ; Nkala Valley, near Lundazi, 
3400 ft, 2 <$, 19.iii.1939 (F. B. Macrae) ; Malawi: Kasangazi, near Bandawe, 3000 ft 
above L. Nyasa, 2 <$ (Prentice) ; Magunda Estate, Luchenza, 1 9 (F- Nisbet) ; Mt. 
Mlanje, 1 6*, 27.iii.1913; 2 <$, 2.^.1913 (5. A. Neave); Zomba Plateau, 1 <$, iv.1920 



44 P. R- SEYMOUR 

(H. Barlow) ; Zomba, i £, vi.1923 (H. Barlow) ; Congo (Kinshasa) : 1 <$, Elisabeth- 
ville, 1 $, iii.1925, 1 $, 27.iii.1934, 1 $ (Bourguignon) , in MRAC, Tervuren; 2 $, 
20.iii.1943 (C. Seydel), in ZSBS, Munich; 1 J, 20.iii.1950 (C. Seydel), in MRAC, 
Tervuren; 9-14.iv.1955 (C. Seydel), in ZSBS, Munich; Katanga, Kafakumba, 7 (J, 
1 $, hi. 1927; Kambove, 4000-5000 ft, 1 <$, 4.^.1907; Kassonyi, 1 <$, viii.1937 
(/. Breolo), in MRAC, Tervuren; Likasi, 4000 ft, 1 <J, 20.iii.1919 (T. A. Barns); East 
Luvua Valley, 5700 ft, 2 <J, ii.1922 (T. A. Barns) ; 3000 ft, 1 <J, iv.1922 (7. A Barns) ; 
Omkoro, 1 (J, 6. hi. 1926; Usumbura, 1 <$, 5JV.1926 (F. G. Jackson); Tanzania: 
Kigonsera, 1 g, in ZSBS, Munich; Kilosa, 1 <$, 19.iv.1923 (Loveridge) ; Madibira, 1 <J, 
in ZSBS, Munich; Marungu Plateau, 7000 ft, 2 <$, 4 $, ii.1922 (T. A. Barns) ; Mbeya, 
1 <J, (E. Ross) ; 1 J, hi. 1950 (iV. Mitton) ; Musoma, Banagi Hill, 3 $, 2 $ (M. S. Moore) ; 
Shinyanga, Mwandui, 4^, ii.1952 (Croft); Ukerewe Island, 1 <$ (Conrad); Rwanda: 
Gabiru, 1 $, 18. x. 1932 (L. A. Burgeon), in MRAC, Tervuren; Uganda: i $ 
(W. L. Doggett) ; 1 <$, 17.hi.1923 (H. Hargreaves) ; Ankole, i<J, 5.vii.i928, i<J, 14.x. 1928 
(both /. Gastrell); Motuba Unyoro, 3 $, 18.X.1901 (C. S. Betton); North Buddu, 
3800 ft, 1 $, 18.ix.1911 (5. A. Neave) ; south of Lake George, 3200 ft, 1 (J, 18.X.1911 
(S. A. Neave); Kickwamba, 1 $ (C. Christy); Karamoja, Mt. Toror, 1 $, vi.1949 
(T. H. E. Jackson); Mbarara, 1 $ (R. E. McConnell); Mulema, 1 $, v.1903 
(W. L. Doggett) ; S. E. Ruwenzori, 3500 ft, 2$, 16 & 20.iv.1906 (G. Legge & A. F. R. 
Wollaston) ; Kenya : Athi River, 1 <$, 10. v. 1899 (C. S. Betton) ; Eburra, 1 <$, 21. hi. 1900, 
1 $, 25.hi.1900 (both C. S. Betton); Mt. Elgon, 1 $, iv.1925 (G. W. Jeffery); 1 $, 
v.1934 (T. H. E. Jackson) ; south-west slopes, 2 (J, 1 $ (H. B. Labbury) ; South 
Kavirondo, Suna, 1 $, 20. hi. 1930; 3 <$, iv.1930; 1 $, ix.1930; 1 <J, x.1930; 18 <$, 
1 $, ih.1931; 5 ct, iv.1931; 1 <J, x.1931; 7 J, 2 $, xi.1931; 11 <$, 1 $, xh.1931; 3 <J, 
i.1932; 1 (J, 2 $, hi. 1932; 14 (J, 12 $, iv.1932; 1 (J, 1 $, vi.1932 (all W. Feather); 
Kibauni, 2 <3\ 1 $, 4-5. xh. 1898 (i?. Crawshay); Kibwezi, 1 $, xh.1921 (W. Feather); 
Kikuyu, near Nairobi, 5400 ft, 1 <$, 24. hi. 1900 (R. Crawshay) ; Kitale, 1 J, 10. iv.1931 
(G. W. Jeffery); 6 <$, vi-vh.1934 (G. W. Jeffery); Lumbwa, 1 $, 8JV.1923 
(G. W. Jeffery); Machakos, 1 J, 13.V.1898; 1 £, 6.VL1898; 1 $, 10.xh.1898 (all 
R. C. Crawshay) ; Mosangaleni, 1 $, 25.lv. 1928 (G. H. E. Hopkins) ; Nairobi, 1 $; 1 <£, 
19.ih.1905 (F. J. Jackson) ; 11 <$, 1 $, iv.1905 (F. /. Jackson) ; 5650 ft, 1 <$, n.v.1915 
(A. Loveridge); 1 $, 25.iv.1916 (W. A. Lamborn); 1 $, 23.vi.1918; 1 <J, x.1918; 1 J, 
xi.1918 (all W. H. van Someren); 2 <$, hi. 1926 (yaw Someren); 1 (J, hi. 1927 
(Z). M. Hopkins) ; 2 J, 1 $, iv.1927 (Z>. M. Hopkins) ; 1 $, xi.1928; 2 ^, iv.1937; 1 J 1 , 
vii-viii.1936 (all van Someren) ; Nakuru, 2 $, 2 $, 1 <$, 6.VL1943; 1 <$, 9.^.1947; 1 <$, 
i.v.1949; 1 $, 18.V.1950; 1 <$, n.v.1952 (all A. Townsend); Nakutu, 1 <$, 8.V.1921 
(H. A. Bodeker) ; Nandi, Moboroni, 1 $, vh.1903 (J. J. Jackson) ; Ngong, 1 <$, vi.1943 
(van Someren); Ethiopia: Dangila, 2 $, 9-10. ix. 1926; 1 <$, 29.vih.1930; 3 <$, 13. ix. 
1930; 1 <J, 1 $, 1 & 26. ix. 1932; Dib Qan, 1 £, 22. ix. 1932 (all R. E. Cheesman) ; Harar, 
1 $, 27.vih.1937; 2 (J, ix-x.1939 (all R. E. Ellison) ; Sudan: Blue Nile Province, Toz, 
1 <$, 20.vih.1960 (H. Schmutterer) , in ZSBS, Munich; Darfur Province, Kulme, 7 $, 
vi-x.1921 (H. Lynes) ; Gondokoro, White Nile, 3^ (W. E. Rymes-Cole) ; 2$ (Dabbene) ; 
Kordofan Province, Kadugli, 1 $, 29.vih.1962 (H. Schmutterer), in ZSBS, Munich; 
Mongalla, White Nile, 5 <$, 20.X.1917 (Yardley); Tambura, 4 <$, 2 $; White Nile, 
io°i2'N, 2 $, 5 $ (Yardley) ; India: southern India, 6 ^, 2 $; Madras, Nilgiris, 1 <$, 



REVISION OF GENUS MASALIA 



45 



I 9: Madhya Pradesh, Jubblepore, i $, 17.ix.1905 (C. S. Betton); M. P., Mhow, 
1 cJ, 1 $, ix.1886 (C. Swinhoe); M. P., Wynaad, 2 (J; Mysore, Belgaum, 1 $ 
{T. R. Bell); M., Belgaum, 1 ?, 29.ii.1922 (T. fi. fie//). 

Distribution (Map 5). Africa south of the Sahara; central and southern India. 

Remarks. This subspecies may be recognized by the modified form of the 
papilla analis, which is simple in M. g. bechuana. 

The synonymy of Timora lancea is based on a photograph presented by Dr E. Berio 
of one of the paratypes, and on a specimen from the BMNH Collection identified by 
him. 




A galatheae galatheae 
A galatheae bechuana 



Map 5. Distribution of subspecies of M. galatheae. 



46 



P. R. SEYMOUR 



Masalia galatheae bechuana subsp. n. 

(Text-figs 50, 51; PI. 3, figs 145-149; Ma P 5) 

Fore wing with areole; length <$ (10), 12 -6-15-0 (holotype 14-2), $ (18), 12-7-16-5. Wing 
pattern as in PI. 3, figs 145-149. Fore wing upper surface with costal and anal regions either 
light yellow or dull red to greyish pink; central region pale yellow immaculate or irrorate 
between veins with dark brown. Hind wing upper surface white to pale yellow. 

Genitalia, <J scobinate bar and cornutus as in Text-fig. 51. $ papilla analis simple (Text- 
fig. 5°)- 

Material examined. 

Holotype, [Botswana] British Bechuanaland : Upington, $, 13.iii.1950 
{H. B. Kettlewell). 

Paratypes. South West Africa: Hoffnung, 1850 m, 3 J, 10. i. 1934; Okahandja, 
1 <?, 10-16.ii.1928 ; 1 $, 2-18. iii. 1928 (both R. E. Turner) ; 1 $, 4.U1.1949 ; 1 $>, 30.1.1953 ; 
1 (J, 1 $, 30 and 25.1.1954; 3 $, 24-25. iii. 1954 (all F. Gaerdes), in ZSBS, Munich; 
Windhoek, 1650 m, 2 $, 16 and 23. i. 1934 {K.Jordan) ; [Botswana] British Bechuana- 
land: Okahandja, 1320 m, 1 <$, 1-4. i. 1934 (K. Jordan) ; Upington, 1 <$, 10 $, 12-14. iii. 
1950 (H. B. Kettlewell); [South Africa] Transvaal: Zutrsenka (Waterberg District), 
1 <$, i-iii.1899. 

Distribution. South West and South Africa. 




..:* 




;; .,-;-/ 



Figs 50-51. M. galatheae bechuana, genitalia. 50, $, papilla analis. 
51, o\ scobinate bar and cornutus. 



REVISION OF GENUS MASALIA 



47 



Remarks. M. g. bechnana can be separated from the nominate subspecies on 
difference in the female papilla analis, simple in M. g. bechuana, of modified form in 
M. g. galatheae. 

Masalia cruentata (Moore) comb. n. 
(Text-figs 52-54; PI. 4, figs 150-152; Map 6) 

Curubasa cruentata Moore, 1881 : 367. LECTOTYPE o*. India (BMNH), here designated 

[examined]. 
Curubasa marginata Moore, 1881 : 367. LECTOTYPE $, India (BMNH), here designated 

[examined]. [Synonymized by Hampson, 1903 : 112.] 
Timora cruentata (Moore) Hampson, 1903 : 112. 




52 






X 

* 



53 




Figs 52-54. M. cruentata, genitalia. 52, $, papilla analis. 53 and 54, $, scobinate 

bar and cornutus. 



48 



P. R. SEYMOUR 



Fore wing with areole usually present; length $ (52), 11 -2-18-5 (lectotype 14-1), $ (32), 
12-7-18-2. Wing pattern as in PI. 4, figs 150-152. Fore wing upper surface with costal 
region white to pale yellow, immaculate or costa tipped with pink or pale or greyish red; 
occasionally entire costal region pink, pale or greyish red. Central region white to pale yellow; 
anal region pink, pale or greyish red or light to reddish brown. Hind wing upper surface white 
to pale yellow, immaculate or lightly irrorate with greyish brown. 

Genitalia, o* scobinate bar and cornutus as in Text-figs 53, 54. $ papilla analis modified, 
terminal spines absent (Text-fig. 52). 

Material examined. 

Curubasa cruentata, LECTOTYPE, here designated, [India:] North West India 
(Cashmir, Noashera — from original description), (labelled 9) ; paralectotype, [India:] 
North West India, $. Curubasa marginata, LECTOTYPE, here designated, 
[India]: North West Himalaya, $; paralectotype, [India:] North West India, 9- 

India: Himachal Pradesh, Dharmsala, 3$; H.P., Kasauli, 1 $, 28.viii.1905; H.P., 
Kula District, 1 <J, 5 9; H.P., Sabathu, 1 $, vii.1889; H.P., Solon, 3 <J, viii.1896; 1 <J, 
ix.; H.P., Sultanpur, 1 9; Nepal: Pakhara, 28°i4'N, 83°59'E, 1 <$, 16. ix. 1955; 
Tuckucha, 28°43'N, 83°39'E, 1 9, 12. ix. 1955 — both Zool. Staatssamml., Miinchen. 
Tibet: Batang, 2800 m, 1 9. 17.viii.1936; 1 $, 21.viii.1936 — both Mus. Bonn; 
China: Chekiang, West Tien-mu-shan, 1 <$, 24.iv.1932; Shantung, Tai-shan, 1550 m, 
1 9, vi.1934; 1 (J, 3.viii.i934; 1 9, 5.VIU.1934; 1 <J, 4.viii.i934; 1 <J, 1 $, 8.viii.i934; 




• cruentata 



Map 6. Distribution of M. cruentata. 



REVISION OF GENUS MASALIA 49 

2 $, 9.viii.i934; 10 <J, 3 $, 11.viii.1934; 5 <J, 1 ?, 12.viii.1934; 5 <$, 4 $, 13.viii.1934; 
icJ, 17.viii.1934; i<J, 1 $, 28.viii.1934 — all in MAK, Bonn; Tsekou, 3 (J, 2 $; Yunnan, 
1 (J, 1 $, 1918; Yunnan, Li-Kiang, 2000 m, 1 $, 24.vii.1934; 1 J, 8.viii.i934; 1 $>, 
18. x. 1934; 1 $, 25.vii.1935 — Li-Kiang material from MAK, Bonn. 

Distribution (Map 6). North India, Nepal, Tibet and south-west and central 
east China. 

Remarks. Differences between M. cruentata and the closely related M . galatheae 
are given under the heading of galatheae-crnentata complex. The material from 
Tai-shan is atypical in that, with few exceptions, an areole is absent; a long series 
was collected from this locality. The similarity to one another in other respects of 
these specimens, and the short period over which they were caught, suggests that 
they formed part of a local inbred population. 



Masalia flaviceps (Hampson) comb. n. 
(Text-figs 55, 56; PI. 4, fig. 159) 
Timora flaviceps Hampson, 1903 : 116. Holotype <$, Nigeria (BMNH) [examined]. 

Fore wing without areole; length <$ (34), no-132 (holotype 12-0), $ (6), 12-4-14-0. Wing 
pattern as in PI. 4, fig. 159. Fore wing upper surface white; in a few specimens the costal and 
anal regions are slightly tinged with yellow, with just sufficient contrast for divisions between 
the regions to be made. Hind wing upper surface white. 

Genitalia. <$ scobinate bar and cornutus as in Text-fig. 56. $ papilla analis modified; 
terminal spines present and dorso-lateral surface clothed in fine hair (Text-fig. 55). 

Material examined. 

Timora flaviceps, holotype, Nigeria: Borgu, Yelwa Lake, <J, 2.x. 1899 (G. Migeod). 

Senegal: Kaolack, 7 <£, 1 $, 1909 (G. Melon); Ghana: Kete-Krachi, 16 <$, 3 $ 
(A W. Cardinall) ; Navaro, 2 J, 1 $, x.1923 (/I. W. Cardinall) ; Nigeria: Minna, 1 <£, 
1 $, 13.X.1910 (Scott Macfie) ; Zungeru, 2 <$, 25.X.1910 (Scott Macfie) ; Sudan: White 
Nile, n°i2'N, 2 cJ (Yardley); n°i5'N, 2 <$ (Yardley); Ethiopia: i £ (paralectotype 
of Timora hololeuca Hampson). 

Distribution. From West Africa to Ethiopia. 

Remarks. M . flaviceps is probably most closely allied to M . galatheae. Struc- 
turally there is no apparent difference, and differentiation of the wing regions trace- 
able in a few specimens of M. flaviceps reveals the M. galatheae pattern. Never 
the less, on colour the two species are readily distinguished : in M . flaviceps the costal 
and anal regions are white or white tinged with yellow, whereas in M. galatheae the 
costal and anal regions are pink. M. hololeuca, the only other white species in 
Masalia, can be separated on difference in the form of the female papilla analis: 
modified in M. flaviceps, simple in M. hololeuca. 



50 



P. R. SEYMOUR 



Masalia hololeuca (Hampson) comb. n. 
(Text-fig. 57; PI. 4, fig. 153) 

Timora hololeuca Hampson, 1903 : 117. LECTOTYPE $, India (BMNH), here designated 
[examined] . 

Fore wing with areole; length o* (10), 11 -1-13-5 (lectotype 11-9), $ (10), 12-6-15-0. Wing 
pattern as in PL 4, fig. 153. Fore wing upper surface white. Hind wing upper surface white. 
Genitalia. $ scobinate bar and cornutus as in Text-fig. 57. $ papilla analis simple. 

Material examined. 

LECTOTYPE, here designated, [India: Mysore] Madras, Belgaum, <$, x.1896 
[Watson). 

India: Maharashtra, Nasik, 2 J, 1 $, ix.1892 ; southern India, 5 $, 5 $ [T. R. Bell) ; 
?, C. Rock, 2<J, 5 $, x. 1892. 

Distribution. Central and southern India. 

Remarks. M. hololeuca can be distinguished from M. flaviceps (the only other 
white species in the genus) on the absence and presence, respectively, of an areole, 




55 




\ 56 



Figs 55-56. M. flaviceps, genitalia. 55, $, papilla analis. 56, <J, scobinate bar 

and cornutus. 



REVISION OF GENUS MASALIA 



5i 



and in the female, on the difference in form of the papilla analis, simple in 
M. hololeuca and modified in M. flaviceps. 

The type-series of this species comprised three males, two from India and one 
from Ethiopia. The Ethiopian male is in fact a specimen of M. flaviceps, and the 
second Indian male is lost. 

Masalia radiata Moore 

(Text-figs 58-61 ; PI. 4, figs 154-158) 

Masalia radiata Moore, 1881 : 364. 

M. radiata, though showing variability in colour, is marked similarly to other species 
within the group. Two subspecies are recognized, the nominate subspecies, a 
small brown moth from northern India and M. r. terracotta, larger, light to reddish 
brown, from central and southern India, west Bengal and Sikkim. Within the 
radiata-gvoup, radiata can be separated from all but one species, flavistrigata, on 
differences in structure of the female genitalia. In radiata the papilla analis is 
modified; in rubristria, beatrix and epimethea, simple. In roseivena and rosacea, 
which have modified papilla anali, terminal spines are present, but these are lacking 
in radiata. Though the relationship between radiata and flavistrigata is probably 
no closer than between radiata and any other species in the group, no single character 
common to both subspecies has been found to separate radiata from flavistrigata. 




Fig. 57. M. hololeuca, q\ scobinate bar and cornutus. 



52 P. R. SEYMOUR 

Each subspecies however, can be separated using a different single character, the 
brown upper surface of the hind wing in M. r. radiata and the non-overlapping 
scobinate bar spicules in M. r. terracotta. In flavistrigata the upper surface hind 
wing colour is white and the scobinate bar spicules overlap. 

Masalia radiata radiata Moore 
(Text-fig. 59; PI. 4, figs 154, 155) 

Masalia radiata Moore, 1881 : 364. LECTOTYPE $, India (BMNH), here designated 
[examined]. 

The subspecies is known only from male specimens. The papilla analis is assumed 
to be as in M. r. terracotta. 

Fore wing with areole; length, <$ (2), io-6 (lectotype). Wing pattern as in PI. 4, figs 154, 155. 
Fore wing upper surface with ground colour brownish orange to light brown; upper central 
longitudinal streak white. Hind wing upper surface brown. 

Genitalia, o* scobinate bar and cornutus as in Text-fig. 59. 

Material examined. 

LECTOTYPE, here designated, [India: Uttar Pradesh,] N. W. Provinces, 
[Mainpuri] Manpuri, 1 <$. 

India: Rajasthan, Ajmer, 1 J, 14.viii.1892. 

Distribution. North India. 

Remarks. Externally M. r. radiata can be distinguished from M. r. terracotta on 
size and colour of hind wing. M. r. radiata is smaller (fore wing length 10 -6 mm in 
radiata, 11 -0-15 -4 mm in terracotta) with brown hind wings; the hind wings of 
M. r. terracotta are white or greyish to brownish orange. The males of the two sub- 
species can be distinguished on the closeness of the scobinate bar spicules: over- 
lapping in M. r. radiata, non-overlapping in M. r. terracotta. 

Masalia radiata terracotta Hampson stat. n. 
(Text-figs 58, 60, 61 ; PI. 4, figs 156-158) 

Masalia terracotta Hampson, 1891 : 71. LECTOTYPE 0*. India (BMNH), here designated 

[examined]. 
Timora flavia Hampson, 1903 : 113. LECTOTYPE £, India (BMNH), here designated 

[examined]. Syn. n. 

Fore wing with or without areole; length $ (32), 11 0-13-5 (lectotype), $ (15), i2-2-i5*4. 
Wing pattern as in PI. 4, figs 156-158. Fore wing upper surface with ground colour pale 
yellow or light to brownish orange. Upper central longitudinal streak white. In specimens 
with pale yellow grounds, the upper central longitudinal streak may be outlined or partly out- 
lined in greyish red or light brown. Hind wing upper surface white or greyish to brownish 
orange. 

Genitalia. <J scobinate bar and cornutus as in Text-fig. 61. $ papilla analis modified, 
terminal spines absent (Text-figs 58, 60). 



REVISION OF GENUS MASALIA 



53 






Figs 58-61. 58, M. radiata terracotta, $, papilla analis. 59, M. r. radiata, £, scobinate bar 
and cornutus. 60, M. r. terracotta, $, papilla analis. 61, M. r. terracotta, $, scobinate 
bar and cornutus. 



54 p. r- seymour 

Material examined. 

Masalia terracotta, LECTOTYPE, here designated, [India: Madras,] Nilgiris, $, 
paralectotypes, [India: Madras,] Nilgiris, i <$, i $. Timor a flavia, LECTOTYPE, 
here designated, [India: Mysore,] Belgaum, <$, x.1896 {Watson); paralectotypes, 
(India: Madhya Pradesh,] Mhow, 1 $, ix.1881 (C. Swinhow); [Mysore,] Belgaum, 
1 $, ix.1896 (Watson); Deccan, 1 J (Sykes); southern India, Wynad, 1 <$. 

India: Maharashtra, Bombay, 3 <J; M., Nasik, 3 $; Mysore, Belgaum, 1^,1$, 
ix.1896; 2 (J, 4 $ (T. R. Bell) ; Kerala, Peermade, 2 <$, 1 $ (Imray) ; K., I <J; southern 
India, 2 6* (7. i?. £^/) ; Madras, Nilgiris, 3000 ft, 1 $ (Hampson) ; Madhya Pradesh, 
Mhow, 1 (J, 2 $, ix.1881 (C. Swinhoe); West Bengal, Darjeeling, 1 $ (Pitcher); 
Sikkim: 1 (J, v.1893. 11 S> 2 $ specimens with localities indistinctly written. 

Distribution. India and Sikkim. 

Remarks. The two subspecies can be separated on difference in size and hind 
wing colour: M. r. terracotta is larger (compare fore wing length) with white or 
greyish to brownish orange hind wings; the hind wings in M. r. radiata are brown. 
Non-overlapping scobinate bar spicules in the male of M. r. terracotta, contrasted 
with the overlapping spicules in M. r. radiata, also distinguish between them. 



Masalia rubristria (Hampson) comb. n. 
(Text-figs 62-64; PL 5, figs 162-170; Map 7) 
Timora rubristria Hampson, 1903 : 112. 

M. rubristria is a variable species both in colour and marking, closely allied to 
M. epimethea and M. beatrix. The species is divided into three subspecies. Of 
these three, two (M, r. rubristria and M. r. transvaalica) have no areole and, on the 
absence of this character, are separated from M. epimethea and M . beatrix. The 
third subspecies, in which an areole is present, M. r. rhodomelaleuca, can be dis- 
tinguished from M. epimethea and M. beatrix on differences in fore wing colour. 

M. rubristria can be separated from the remaining species in the ra<&ata-group 
(the only others in the genus likely to be confused with it) by the form of the female 
papilla analis, simple in M. rubristria, modified in M. flavistrigata, M. radiata, 
M. rosacea and M. roseivena. 

Differences occur between monomorphs from a number of localities ; the similarity 
of monomorphs from others, and from at least one locality the occurrence together 
of dimorphs, each variant known from at least one other locality as a monomorph 
(see under discussion of M. r. rubristria), cannot be satisfactorily resolved without 
further material and investigation. The tentative division of the material into three 
subspecies is primarily based on geographic separation of the known material. 
Structurally however, on the presence of an areole, M. r. rhodomelaleuca can be 
separated from subspecies rubristria and transvaalica. M. r. rubistria and M. r. 
transvaalica differ in the colour of their fore wing markings, light brown, brownish 



REVISION OF GENUS MASALIA 55 

or reddish brown in nibristria, pale to pastel or greyish red in transvaalica. Speci- 
mens which are not clearly recognized on colour may be separated on size; of the 
two, transvaalica is larger though the ranges of the two subspecies do overlap. 



Masalia rubristria rubristria (Hampson) 
(Text-figs 62, 63; PL 5, figs 162-165; Map 7) 

Timora rubristria Hampson, 1903 : 112. Holotype £, Nigeria (BMNH) [examined]. 
Timora multistriata Bethune-Baker, 191 1 : 507. LECTOTYPE <J, Nigeria (BMNH), here 

designated [examined]. Syn. n. 
Timora sanguistria Berio, 1966 : m. Holotype <J, Senegal (MNHN, Paris) [examined]. 

Syn. n. 

Fore wing without areole; length^ (32), 11-4-15-3 (holotype 11-9), $ (20), 11-9-16-2. Wing 
pattern as in PI. 5, figs 162-165. Fore wing upper surface with pattern and colouring variablein ; 
PI. 5, figs 162, 163, 164, central and anal regions white with white-brown, brown, or reddish 
brown markings; costal region either yellowish white, pale yellow, light brown or brown; in PI. 5, 
fig. 165, ground colour pale yellow, upper central and anal longitudinal streaks white, outlined 
with greyish rose. Hind wing upper surface white, marginally irrorate with light brown, or 
light brown (sanguistria holotype). 

Genitalia, o* scobinate bar and cornutus as in Text-figs 62, 63. $ papilla analis simple. 

Material examined. 

Timora rubristria, holotype, Nigeria: Asaba, $ (Crosse). Timora multistriata, 
LECTOTYPE, here designated, Nigeria: 100 miles north of Lokoja, J (D. Cator). 
Timora sanguistria, Holotype, Senegal: Badi, <$ [Institut francais d'Afrique noire 
Dakar (Senegal)], MNHN, Paris. 

Senegal: Kaolack, 1 $ (G. du Dresnay), in MNHN, Paris; 1 $ (LeMoult) ; Guinea: 
Diondougou, 1 <$ (L. J. De Joannis), in MNHN, Paris; Ghana: Gambaga, 11 <J, 11 $ 
(Bury); Kete-Krachi, 6 <$, 5 $ (A. W. Cardinall); [Nararo] Navaro, 2 <J 
(A.W.Cardinall) ; Nigeria: Ogruga [POgrugru], River Niger, 1 $; Baro, 1 $, 25.ix.1910 
(Scott Macfie); Minna, 2 $, i.x.1910 (Scott Macfie); 100 miles north of Lokoja, 1 <$ 
(D. Cator); Central African Republic: Fort Crampel, 3 <$, 2 $ (LeMoult). 

Distribution (Map 7). Senegal, Guinea, Ghana, Nigeria and Central African 
Republic. 

Remarks. The absence of an areole and the brown or reddish brown fore wing 
markings in M. r. rubristria separate it from M. r. rhodomelaleuca (areole present) 
and M. r. transvaalica (fore wing pale, pastel-, or greyish red), respectively. 

Specimens similar to the lightly marked one from Guinea are known from Minna 
(Nigeria) and Nararo (Ghana) (Text-fig. 165). A second variant (Text-fig. 162) is 
also known from this latter locality. Specimens from Fort Crampel (Central African 
Republic), Kete Krachi (Ghana) and Gambaga (Ghana) are essentially alike. Fort 
Crampel specimens are larger and have slightly wider, white longitudinal streaks. 
Of the remaining material variation occurs from locality to locality. 



50 



P. R. SEYMOUR 





Figs 62-64. M. rubristria subspecies, $, scobinate bar and cornutus. 62 and 63, 
M. r. rubristria. 64, M. r. transvaalica. 



REVISION OF GENUS MASALIA 



57 



Masalia rubristria transvaalica (Distant) comb, et stat. n. 

(Text-fig. 64; PI. 5, figs 166-169; Map 7) 

Timora transvaalica Distant, 1902 : 212. LECTOTYPE $, South Africa (BMNH), here 

designated [examined]. 
Timora rosea Gaede, 1915 : 39. LECTOTYPE <$, South Africa (MNHU, Berlin), here 

designated [examined]. Syn. n. 

Fore wing without areole; length^ (21), 13-8-17-5 (lectotype 17-3), ? (4), 15-8-17-8. Wing 
pattern as in PI. 5, figs 166-169. Fore wing upper surface with pattern and colouring variable: 
in PI. 5, figs 166, 167, ground colour pale to pastel-red, occasionally with a number of veins 
lightly outlined in brown, upper central and anal longitudinal streaks (when present) white ; in 
PI. 5, figs 168, 169, ground colour yellowish white to pale yellow, radial, medial, cubital and anal 
veins outlined in pastel to greyish red. Hind wing upper surface white, pale to pastel-red, or 
light brown to brown. 

Genitalia. $ scobinate bar and cornutus as in Text-fig. 64. ? papilla analis simple. 





k\ 




^P rubristria rubristria 
A rubristria transvaalica 



rubristria rhodomelaleuca 



Map 7. Distribution of subspecies of M. rubristria. 



58 p. r. seymour 

Material examined. 

Timor a transvaalica, LECTOTYPE, here designated, [South Africa :] Transvaal, 
Johannesburg, $ (J. Hyde) ; Timora rosea, LECTOTYPE, here designated, [South 
Africa:] Transvaal, White River, <$, ii.1911 {A. T. Cooke), in MNHU, Berlin. 

Kenya: Kavirondo, Suna, 10 g, iii.1931; 3 $, iv.1931; 1 <$, xi.1931; 1 J, ii.1932; 
Tanzania: Kalambo River, 1 $; Kilossa, 1 <J, 28.iii.1922; Njombe, 6000-6500 ft, 
1 (J, 19. ii. 1952 (W. Peters); Congo (Kinshasa): Elisabethville, 2 <$, 3, 5.111.1949 
(Ch. Seydel), in MRAC, Tervuren; Zambia: Kapapi, 1 <J, 17.iii.1939 (F. B. Macrae); 
Rhodesia: Sinoia, 1 <J, 21. ii. 1950 (N. Mitton); South Africa: Transvaal White 
River, 1 <$, io.i.1910 (A. T. Cook) ; 2 $ (C. H. Pead). 

Distribution (Map 7). Kenya, Tanzania, Congo (Kinshasa), Zambia, Rhodesia 
and South Africa. 

Remarks. The absence of an areole from M . r. transvaalica separates it from 
M. r. rhodomelaleuca; M. r. transvaalica and M. r. rubristria are fairly readily 
distinguished on the colour difference of the fore wing markings, pale, pastel to 
greyish red in M. r. transvaalica, brown or red in M. r. rubristria. 

Across the range of M. r. transvaalica differences in colour and pattern occur in a 
number of localities. 

Masalia rubristria rhodomelaleuca (Berio) comb, et stat. n. 

(PI. 5, fig. 170; Map 7) 

Timora rhodomelaleuca Berio, 1943 : 182. Holotype o*. Ethiopia: Elaberet, 17. ix. 1938 
(G. Vaccaro) (Berio Coll., MCSN, Genoa). 

Fore wing with areole; length $ (1), 15-4. Wing pattern as in PI. 5, fig. 170. Fore wing 
upper surface with costal and anal regions light to greyish orange ; central region with proximal 
and disto-radial areas brown, disto-cubital area light to greyish orange; upper central longi- 
tudinal streak white. Hind wing upper surface light to yellowish brown. 

Genitalia. $ papilla analis simple. 

Material examined. 

[Paratype] allotype, [Ethiopia] Eritrea: Elaberet, $, i.ix.1938 (F. Vaccaro), in 
MCSN, Genoa. 

Distribution (Map 7). Ethiopia. 

Remarks. The presence of the areole in M. r. rhodomelaleuca distinguishes it 
from both M. r. transvaalica and the nominate subspecies. 

Masalia beatrix (Moore) comb. n. 

(Text-figs 65-66; PL 6, figs 174-178) 
Pradatta beatrix Moore, 1881 : 365. 
Fore wing with areole. 

Two subspecies are recognized, beatrix from India and trifasciata from Africa. 
Whilst the fore wing pattern in the African subspecies is fairly constant, that of the 



REVISION OF GENUS MASALIA 59 

Indian subspecies is variable and makes separation of the two difficult. The fore 
wing central region in M. b. trifasciata differs in colour from the costal and anal 
regions and is clearly deliminated from them; there is always a well differentiated 
brown to dark brown lower central longitudinal streak. In M. b. beatrix the central 
region is similarly coloured to the costal and anal regions and with incursion of 
colour from these outer regions, the central region is not clearly deliminated. The 
lower central longitudinal streak, when present, ranges from light orange to pastel- 
red. Occasionally it is suffused with brown but in contrast to the brown streak in 
M. b. trifasciata, any brown marking present is narrow and poorly defined. 

M . beatrix is distinguishable from other species in the radiata-group on genitalic 
characters. The simple form of the papilla analis in M. beatrix separates it from 
M. flavistrigata, M. radiata, M. rosacea and M. roseivena; in the latter four, the 
papilla analis is modified. Overlapping scobinate bar spicules in M. beatrix separate 
it from the remaining two species, the African M. rubristria and the Madagascan 
M. epimethea; in M. rubristria and M. epimethea the spicules are non-overlapping. 

Masalia beatrix beatrix (Moore) 
(Text-fig. 65; PI. 6, figs 174-176) 

Pradatta beatrix Moore, 1881 : 365. LECTOTYPE <J, India (BMNH), here designated 

[examined] . 
Timora beatrix (Moore) Hampson, 1903 : 114. 
Timora nigristriata Hampson, 1903 : no. Holotype <$, India (BMNH) [examined]. Syn. n. 

Fore wing, length o* (12), 12-9-14-9 (lectotype 14-2), $ (17), 12-6-15-4. Wing pattern as in 
PI. 6, figs 174-176. Fore wing upper surface with costal and anal regions orange-white to light 
orange or reddish white to pastel-red, central region white to yellowish white, streaked with the 
same colour as the costal and anal regions and with costal and anal region colouring extending 
in from these regions. Depending upon incursion of colour and extent of streaking a lower, 
central, longitudinal streak may be differentiated; when present the streak is immaculate or 
irrorate with light brown. 

Genitalia, o" scobinate bar and cornutus as in Text-fig. 65. $ papilla analis simple. 

Material examined. 

Pradatta beatrix, LECTOTYPE, here designated, [India: Punjab,] Dharmsala, 
cJ {Hocking); paralectotypes, [India: Punjab,] Dharmsala, 2 $, 2 $ (Hocking); 
Timora nigristriata, holotype [India: Mysore,] Belgaum, <$, ix.1896 (E. Watson). 

India: Rajasthan, Ajmer, 1 <$, 28.vii.1892; 1 <$, 4.viii.i8g2; 1 $, 10.viii.1992; 
Himachal Pradesh, Mandi, 3-5000 ft, 1 $, 1883 (G. Young); H. P., Kangra, 1 $, 
1884; N.W. Himalaya, 1 <$; Maharashtra, Nasik, 1 <£, 4 $, ix.1892; M., Nagpur, 
Chota, 1 $, 1911; Kerala, Peermade, 1 $ (Imray); K., 1 J, 1 $ (Place); southern 
India, 1 $ (T. R. Bell); Madras, Cuddapah, 1 <J (Campbell); M., Nilgiri Hills, 1 $; 
?, Simla, 7000 ft, 1 $ (A. E. Jones); West Bengal, Darjeeling, 1 $ (Pitcher); 1 $ 
(C. R. Oakes). 

Distribution. India. 

Remarks. The absence of a brown lower central longitudinal streak (or at most 



6o 



P. R. SEYMOUR 



presence of a streak irrorate with brown and forming a fine brown streak within the 
longitudinal streak) distinguishes M. b. beatrix from M. b. trifasciata. In M. b. 
trifasciata the lower central longitudinal streak is always well defined and brown to 
dark brown in colour. 



Masalia beatrix trifasciata (Hampson) comb, et stat. n. 
(Text-fig. 66; PI. 6, figs 177, 178) 
Timora trifasciata Hampson, 1903 : no. Holotype $, Kenya (BMNH) [examined]. 

Fore wing, length^ (10), 14-2-15-8, $ (15), 15-0 (holotype)-i6-4 (5). Wing pattern as in PI. 6, 
figs 177, 178. Fore wing upper surface with costal and anal regions from pastel to greyish red; 
anal region lightly to moderately irrorate with dark brown. Central region white to orange- 
white. Lower central, and longitudinal streak between R5-M1, dark brown; R5-M1 streak 
faintly to well developed. Hind wing upper surface white to orange-white, distally light to 
moderately irrorate with brown. 

Genitalia, o* scobinate bar and cornutus (Text-fig. 66). $ papilla analis simple. 

Material examined. 

Holotype [Kenya] B. E. Africa: [Stony Athi, River Athi,] Stony River, $, 
2.XU.1898 (R. Crawshay). 

Kenya: Kavirondo, 1^,1$, iii.1931 (W. Feather); 1 <$, iv.1932 (W. Feather); 
Kibwezi, 1 <$, iii.1922 (W. Feather); Tanzania: Musoma, Banagi Hill, 1 <$, 1 £ 
(M. 5. Moore); Shinyanga, Mwandui, 1 $, xii.1951; 1 <$, i.1952; 1^,1$, ii-iii.1952; 
1 (J, hi. 1952 (all Croft). 

Distribution. Kenya and Tanzania. 







Figs 65-66. M. beatrix subspecies, q\ scobinate bar and cornutus. 65, M. b. beatrix. 

66, M. b. trifasciata. 



REVISION OF GENUS MASALIA 



61 



Remarks. The presence of a pronounced brown lower longitudinal streak in 
M. b. trifasciata distinguishes it from M. b. beatrix. When present, the streak in 
M. b. beatrix is of orange or red colour, occasionally irrorate with brown. A brown 
streak may be formed through dense irroration but in those specimens in which this 
occurs, the streak is always set within a wider streak of orange or red; in 
M. b. trifasciata the central longitudinal streak is unicolorous. 

Masalia epimethea (Viette) comb. n. 



(Text-fig. 67; PI. 4, figs 160, 161) 



Timora epimethea Viette, 1958 
sao, Ambalavao, iii.1956 (A 



: 149. Holotype ?, Madagascar: centre, district de Fianarant- 
Robinson) (MNHN, Paris). 



Fore wing with areole; length <$ (9), 13-0- 15-0, 9 (10), 14-2-157. Wing pattern as in PI. 4, 
figs 160, 161. Fore wing upper surface with ground colour brownish orange, costal margin and 
upper central longitudinal streak white. In a few specimens there is a white longitudinal streak 
between Mz~Cu\ a . When present, terminal and medial dots brown. Hind wing upper surface 
with proximal area either white or light brown; when white, the distal area is light brown; 
when light brown, the distal area is brown; marginal 'cilia' white. 

Genitalia, q* scobinate bar and cornutus as in Text-fig. 67. ? papilla analis simple. 

Material examined. 

Madagascar: i <$, 9 $; de Fianarantsae, 1 <£, iii.1956 (A. Robinson), MNHN, 




Fig. 67. M. epimethea, q", scobinate bar and cornutus. 



62 P. R. SEYMOUR 

Paris (this specimen was collected with the holotype but did not form part of the 
type-series — communication by Dr P. Viette) ; Mananjary, 7 £, 1 2, 1918 (G. Melou). 

Distribution. Madagascar. 

Remarks. Separation of epimethea from other species in the radiata-group 
involves a number of structural characters. The simple form of the papilla analis 
in epimethea separates it from flavistrigata, radiata, roseivena and rosacea; in the 
latter four, the papilla analis is modified. Non-overlapping scobinate bar spicules 
in epimethea separate it from beatrix, in which the spicules overlap ; and the presence 
of an areole in epimethea distinguishes it from rubristria. 



Masalia rosacea Hampson 
(Text-figs 68, 69; PI. 6, figs 179, 180) 

Masalia rosacea Hampson, 1891 : 71. LECTOTYPE q\ India (BMNH), here designated 

[examined] . 
[Pradatta beatrix Moore; Hampson, 1903 : 114. M. rosacea considered to be an aberration of 

P. beatrix Moore.] 
Timora rosacea (Hampson) Warren, 1913 : 315. [Reinstated as valid name.] 

Fore wing with areole; length q* (7), 13-4 (lectotype)-i5-9, $ (2), i6-i-i6-5. Wing pattern as 
in PI. 6, figs 179, 180. Fore wing upper surface with ground colour pale to pastel-red, upper 
central longitudinal streak white to yellowish white. Hind wing upper surface reddish golden 
brownish orange. 

Genitalia. $ scobinate bar and cornutus as in Text-fig. 69. $ papilla analis modified, 
terminal spines present (Text-fig. 68). 

Material examined. 

LECTOTYPE, here designated, [India: Madras,] Nilgiris, <$ {Hampson); para- 
lectotypes, [India: Madras,] Nilgiris, 3 <$ {Hampson). 

India: Kerala, 1 $ {Place) ; K., Peermade, 1 <$, 1 2 {Imray) ; Madras, Nilgiris, 1 <J, 
7.1X.1903 {Hampson); ?, Seven Valley, 1 2. 

Distribution. Southern India. 

Remarks. Within the radiata-group rosacea is separable from all but roseivena 
on features of the papilla analis. The modified form of the papilla analis in rosacea 
separates it from beatrix, epimethea and rubristria (simple in these three) ; and the 
presence of terminal spines separates it from flavistrigata and radiata ; in flavistrigata 
and radiata, terminal spines are absent. M. rosacea is distinguishable from roseivena 
on the difference in the upper surface hind wing colour, reddish golden brownish 
orange in rosacea, white either immaculate or lightly suffused with reddish brown in 
roseivena. 



REVISION OF GENUS MASALIA 



63 



Masalia roseivena (Walker) comb. n. 
(Text-figs. 70, 71; PI. 6, figs 181-184) 

Leucania roseivena Walker, 1866 : 1954. Holotype <$, Flores (UM, Oxford) [examined]. 

Timora roseivena (Walker) Warren, 191 3 : 314. 

Leucania alarioides Butler, 1886 : 392. Lectotype <J, Australia (BMXH) [examined.] 

[Synonymized by Warren, 1913 : 314.] 
Timora alarioides (Butler) Hampson, 1903 : m. [Lectotype designated.] 

Fore wing with areole present; length rj (8), 12-2-14-8 (holotype), $ (10), 13-9-15-4. Wing 
pattern as in PI. 6, figs 181-184. Fore wing upper surface with costal and anal regions pink, 
central region white; lower central longitudinal streak pink, either immaculate or suffused with 
reddish brown. Hind wing upper surface white, immaculate or lightly suffused with reddish 
brown. 

Genitalia. $ scobinate bar and cornutus as in Text-fig. 71. $ papilla analis modified, 
terminal spines present (Text-fig. 70). 

Material examined. 

Leucania roseivena, holotype, Flores: <$ {Wallace), in UM Oxford; Leucania 





\ 




Figs 68-69. M. rosacea, genitalia. 68, $, papilla analis. 69, <$, scobinate bar 

and cornutus. 



64 



P. R. SEYMOUR 



alarioides, lectotype [Australia:] Queensland, Peak Downs, <$; paralectotype 
[Australia : Queensland,] Gayndah, $. 

Lombok: Pringabaja, i $, iv.1896 (H . Frashstorfer) ; Australia: Northern Terri- 
tories, Darwin, 2 $ (/. S. Litchfield) ; Eureke, 2 <$, 4 $, ii.1903 (Tunney) ; Queensland, 
Geraldtown, near Cairns, 1 <J (Meek) ; Mareeba, 1 (J, 25.xii.1961 (i7. Demarz), in 
ZSBS, Munich; Peak Downs, 1 ^; Townsville, 1 $ (Dodd); [?,] 2 $ (Barnard). 

Distribution. Lombok, Flores and Australia. 

Remarks. Apart from rosacea, roseivena can be separated from all other species 
in the radiata-gvonp on characters of the female genitalia, the modified form of the 
papilla analis and presence of terminal spines. In beatrix, epimethea and rubristria 
the papilla anali are simple in form ; flavistrigata and radiata both have modified 
papilla anali but terminal spines are absent. M. roseivena is distinguishable from 
rosacea on the colour difference of the hind wing upper surface: white, either im- 
maculate or lightly suffused with reddish brown in roseivena, reddish golden brownish 
orange in rosacea. 




\\ \ 




Figs 70-71. M. roseivena, genitalia. 70, $, papilla analis. 71, <J, scobinate bar 

and cornutus. 



REVISION OF GENUS MASALIA 



65 



Masalia flavistrigata (Hampson) comb. n. 

(Text-figs 72, 73; PI. 7, figs 186-188) 

Titnora flavistrigata Hampson, 1003 : 114. Holotype _>, Kenya (BMNH) [examined]. 
Timora lineata de Joannis, 1910 : 224. LECTOTYPE J, Guinka (MNIIN, Paris), here desig- 
nated [examined' 1 . Syn. n. 

Prothoracic tibia with outer claw short or absent. Fore wing with areole; length 5 (168) 
11 -4-16-5 (lectotype 134), ? (29), 12-4-18-2. Wing pattern as in PI. 7, figs 186-188. Fore 
wing upper surface with ground colour ranging from pale to greyish orange, or pastel-red 
through brownish orange to dull red. Upper central longitudinal streak, when present, 
yellowish white. Anal region immaculate or lightly suffused with greyish brown. Hind wing 
upper surface white to yellowish white. 

Genitalia. ^ scobinate bar and cornutus as in Text-fig. 73. papilla ana lis modified, dorso 
lateral surface clothe 1 in tine hair, terminal spines absent (Text-fig. 72). 

Material examined. 

Timora flavistrigata, holotype [Kenya:, Nairobi Plains, Kikuyu, o {R- Crawshay). 
Timora lineata, LECTOTYPE, here designated, [Guinea] H. Guinea: Siguiri, 
Oudoula, (J, in MNHN, Paris. 





71 



Figs 72-73. M. flavistrigata, genitalia. 72, J, papilla analis. 73, #, scobinate bar 

and cornutus. 



66 P. R. SEYMOUR 

Ghana: Northern Territories, Kete-Krachi, 9 <$, 6 $ (A. W. Cardinall); Nigeria: 
Minna, 1 $, 6.X.1910 [Scott-Mac fie); Central African Republic: Fort Crampel, 
1 $ (LeMoult); Angola: Luimbale, Mt. Moco, 1800-1900 m, 122 <$, 14 $, 15-24. iii. 
1934 (K. Jordan); Lunda, Xa-Sengue, 2 <$, 9.iv.ig37 (M. A. Exwell); Quicolungo, 
120 km north of Lucala, 2 $, iv.1936 (R. Braun); Ethiopia: Harar, 1 <$, viii.1939 
(R. E. Ellison) ; Uganda: Mulema, 2 $, 1 $, v. 1903 (W. L. Doggett) ; S. E. Ruwenzori, 
3500 ft, 1 <J, 16.iv.1906 (G. Legge & A. E. R. Wollaston); Kenya: Kitale, 146*, 2 $, 
vi-vii.1934 (G. W. Jeffery); Lumba, 1 <$, 2.iv.ig23 (G. W. Jeffery); Mt. Elgon, 1 1 
iv.1932 (T. H. E. Jackson) ; Tanzania: Dodoma, 1 $, iii. 1950 (N. Mitton) ; Marungu 
Plateau, 7000 ft, 1 £, ii.1922 (T. A. Barns); Malawi: Mt. Mlanje, 1 <$, 24.iv.1923 
(S. A. Neave); Zambia: Abercorn, 1 $, ii. iii. 1954 (D. Vesey '-Fitzgerald). 

Distribution. Guinea, Ghana, Nigeria, Central African Republic, Angola, 
Ethiopia, Uganda, Kenya, Tanzania, Malawi, Zambia, South Africa. 

Remarks. Within the radiata-group fiavistrigata can be separated from all but 
radiata on differences in the female genitalia. The modified papilla analis in 
fiavistrigata separates it from beatrix, epimethea and rubristria, in which the papilla 
analis is simple ; the absence of terminal spines in fiavistrigata separates it from 
roseivena and rosacea, both of which possess terminal spines. Although the relation- 
ship between fiavistrigata and radiata is probably no closer than their relationship to 
any other species in the group, no single character has been found to separate 
fiavistrigata from radiata. The white upper surface of the hind wing and the over- 
lapping scobinate bar spicules in fiavistrigata can, however, be used to separate 
fiavistriata from each of the subspecies of radiata : in M . r. radiata the upper surface 
hind wing colour is brown, and in M . r. terracotta the scobinate bar spicules do not t 
overlap. 

THE latinigra-cheesmanae complex 

Affinities with other species in the genus are not particularly close. M. latinigra 
and M. cheesmanae can be separated from all but one other species, M. fiavistrigata, 
by the presence of a single prothoracic claw ; latinigra and cheesmanae are separable 
from fiavistrigata by the more rounded fore wing apex (Text-figs 23 fiavistrigata, 75) 
and the more elongate scobinate bar (Text-figs 7 3 fiavistrigata, 73, 78, 80, 81). 

Two closely related species are recognized, each with two subspecies. The 
species can be separated on differences in the male antennal segments, lamellate in 
latinigra, simple in cheesmanae. On colour and pattern the nominate subspecies 
are readily distinguished from one another but differences between subspecies 
M. latinigra dangilensis and M. cheesmanae tamburensis are less marked. 

Masalia latinigra (Hampson) comb. n. 
(Text -figs 74-78; PI. 7, figs 189-191; Map 8) 

Timor a latinigra Hampson, 1907 : 243. 

Antennal flagellar segments show marked sexual dimorphism : q* segments lamellate (atypical 
for genus), $ segments simple (Text-figs 76, 77). Prothoracic tibia with one inner claw (Text- 
fig. 74). Fore wing with areole of large size (Text-fig. 75). 

Genitalia, (J scobinate bar markedly elongate (Text-fig. 78). $ papilla analis simple. 

The lamellate antennae of the male are diagnostic. 



REVISION OF GENUS MASALIA 



67 




74 





76 




dorsal 




78 \ 



Figs 74-78. M.latinigra. 74, prothoracic tibia. 75, fore wing venation. 76, ?. antennal 
segments, lateral view. 77 , 6\ antennal segments, lateral view. 78. M. I. latinigra, 
o\ scobinate bar and cornutus. 



P. R. SEYMOUR 






Masalia latinigra latinigra (Hampson) 
(Text-fig. 78; PI. 7, figs 189, 190; Map 8) 

Timora latinigra Hampson, 1907 : 243. LECTOTYPE <$, Uganda (BMNH), here designated 
[examined]. 

Fore wing length 6* (30), 12-9-15-8 (lectotype 14-1), ? (7), 13-2-15-3. Wing pattern as in 
PI. 7, figs 189, 190. Fore wing upper surface with ground colour reddish yellow to light orange, 
lower central longitudinal streak, postmedial and terminal dots, dark brown to black. Hind 
wing upper surface yellowish white, finely to moderately irrorate with dark brown. 

Material examined. 

LECTOTYPE, here designated, Uganda, $ (W. L. Doggett) ; paralectotypes, 
Uganda, 16 <J, 4 $ (W. L. Doggett). 

Uganda: Ankole, 2 $, 29.iii.-26.ix.1929 (/. Gastrell); Elburgin Railway Station, 
2<£, 2.vii.i903; Uganda-Kenya: Mt. Elgon, 1 $, v.1934 (T. H. E. Jackson) ; Kenya: 




W cheesmanae cheesmanae 
ML cheesmanae tamburensis 
© latinigra latinigra 

latinigra dangilensis 



Map 8. Distribution of species and subspecies of the latinigra-cheesmanae complex. 



REVISION OF GENUS MASALIA 69 

Kitali, 3 <$, 1 $, 9-24.iv.1927; Nakuru, 1 $, 16.iv.1941; 3 <J, 2 $, 5-22.iv.1942; I $, 
ii.v.1952; 1 $, 5. v. 1954 (Nakuru specimens, ^4. Townsend). 

Distribution (Map 8). Uganda and Kenya. 

Remarks. The nominate subspecies is characterized by a well developed, broad, 
dark brown to black, lower central longitudinal streak ; absent or poorly differentiated 
in M. I. dangilensis. 

Masalia latinigra dangilensis subsp. n. 

(PI. 7, fig. 191 ; Map 8) 

Fore wing length^ (15), 13-0— 14-8 (holotype 13-8). Wing pattern as in PI. 7, fig. 191. Fore 
wing upper surface with ground colour brownish orange, postmedial and terminal dots brown to 
dark brown; a brown, indistinctly differentiated, lower central longitudinal streak may be 
present. Hind wing upper surface brown. 

Material examined. 

Holotype, Ethiopia: Dangila, 40 mis S. of Lake Tana, 6700 ft, J, 31.viii.1926 
(R. E. Cheesman). 
Paratypes. Ethiopia: Dangila, 6700 ft, 14 (J. 14-29. viii. 1926 (R. E. Cheesman). 
Distribution (Map 8). Ethiopia. 

Remarks. The absence of a lower central longitudinal streak (or at the most the 
presence of a poorly differentiated one) distinguishes M. I. dangilensis from 
M. I. latinigra. 

Masalia cheesmanae sp. n. 

(Text-figs 79-81; PI. 7, figs 192-194; Map 8) 

Antenna with $ and $ flagellar segments simple. Prothoracic tibia with one inner modified 
spine. Fore wing with areole. 

M . cheesmanae can be separated from M. latinigra on the difference in the male 
antennal segments, simple in M. cheesmanae (Text-fig. 79), lamellate in M. latinigra 
(Text-fig. 77). 

This species is named after the late Miss L. Evelyn Cheesman, who collected much 
insect material for the BMNH. 

Masalia cheesmanae cheesmanae subsp. n. 

(Text-fig. 80; PI. 7, figs 192, 193; Map 8) 

Fore wing length <$ (5), 12 2-14 4 (holotype 14-3). Wing pattern as in PL 7, figs 192-193. 
Fore wing upper surface with ground colour orange, costa white, markings brown. Hind wing 
upper surface yellowish white. 

Genitalia. £ scobinate bar as in Text-fig. 80. 

Material examined. 

Holotype, [Congo (Kinshasa) :] Upper Uelle District, Dungu, £, v. 

Paratypes. Senegal: St. Louis, 1 <J, iii.1932, in MNHN, Paris; Cameroun: 



70 P. R. SEYMOUR 

Genderu Mountains, 2600 ft, 1 $, ix.1921 ; [Congo (Kinshasa) :] Upper Uelle District, 
Dungu, 2 6*, ix. 

Distribution (Map 8). Senegal, Cameroun and Congo (Kinshasa). 

Remarks. M. c. cheesmanae can be separated from M. c. tamburensis on the 
difference in upper surface hind wing colour, yellowish white in M. c. cheesmanae, 
brownish orange in M. c. tamburensis. 

Masalia cheesmanae tamburensis subsp. n. 

(Text-fig. 81; PI. 7, fig. 194; Map 8) 

Fore wing length, $ (1), 14-3 (holotype). Wing pattern as in PI. 7, fig. 194. Fore wing 
upper surface with ground colour light orange, costa white, markings brown. Hind wing 
upper surface brownish orange. 

Genitalia. <J scobinate bar as in Text-fig. 81. 

Material examined. 

Holotype, Sudan: [Tambura] Tembura, o*, viii [illegibly written]. 

Distribution (Map 8). Southern Sudan. 

Remarks. M. c. tamburensis can be separated from the nominate subspecies on 
the difference in colour of the upper surface of the hind wing, brownish orange in 
M. c. tamburensis, yellowish white in M. c. cheesmanae. 

Masalia uncta (Swinhoe) comb. n. 
(Text-fig. 82 ; PI. 7, fig. 195) 

Adisura uncta Swinhoe, 1885 : 449. LECTOTYPE q\ India (BMNH) [examined]. 
Timora uncta (Swinhoe) Hampson, 1903 : 109. 

The species is known only from male specimens, and of the two referred to by 
Hampson, only one, labelled type, remains. 

Fore wing with areole absent; length q* (i), 127 (lectotype). Wing pattern as in PI. 7, fig. 195. 
Fore wing upper surface with ground colour light yellow; postmedial streaks on veins Mi to 
C«i a brown; marginal cilia pale red, tipped with white. Hind wing upper surface brownish 
orange to light brown, streaked distally with light yellow; marginal cilia pale yellow. 

Genitalia, a* scobinate bar and cornutus as in Text-fig. 82. 

Material examined. 

LECTOTYPE, here designated, [India: Maharashtra,] Bombay, J, x.1886 
(C. Swinhoe) . 

Distribution. South-west India. 

Remarks. The postmedial streaks against the light yellow ground of the fore 
wing, together with the absence of any other marking, distinguishes the species from 
others in the genus. On wing shape it is probably fairly closely allied to M. albicilia, 
and bears a marked resemblance to species within the genus Adisura. 



REVISION OF GENUS MASALIA 



7* 



dorsal 







I! 




Figs 79-81. M. cheesmanae. 79, <$, antennal segments, lateral view. 80, M. c. cheesmanae, 
o\ scobinate bar and cornutus. 81, M. c. tamburensis , q\ scobinate bar and cornutus. 




Fig. 82. M. uncta, $, scobinate bar and cornutus. 



72 



P. R. SEYMOUR 



Masalia albicilia (Hampson) comb. n. 
(Text-fig. 83 ; PI. 7, figs 196, 197) 
Timora albicilia Hampson, 1903 : 115. Holotype <$, Sikkim (BMNH) [examined]. 

Fore wing without areole; length <$ (4), n-8-12-2 (holotype 12-0), $ (2), 12-7-13-2. Wing 
pattern as in PI. 7, figs 196, 197. Fore wing upper surface variable, light yellow suffused with 
brownish orange; outer margin brownish orange (PI. 7, fig. 196), or brownish orange indistinctly 
marked with light yellow; outer margin brownish orange (PI. 7, fig. 197). Marginal cilia white. 
Hind wing upper surface light brown, marginal cilia pale yellow. 

Genitalia, q* scobinate bar and cornutus as in Text-fig. 83. $ papilla analis simple. 

Material examined. 

Holotype, Sikkim: $, 3.x. 1896 {G. C. Dudgeon). 

India: West Bengal, Darjeeling, Gopaldhara, 1 <$; W.B., Darjeeling, Gopaldhara, 
4720 ft, 1 (J, 1 <j>, ix.1916; W.B., Darjeeling, Gopaldhara, 1 $, 18.ix.1916 (all 
H. Stevens) ; W.B., Darjeeling, 1 $. 

Distribution. North-east India and Sikkim. 

Remarks. The brownish orange colouring and virtual absence of pattern from 
the fore wing upper surface, together with the light brown colouring of the hind 
wing, distinguishes this species from others in the genus. 




83 



Fig. 83. M. albicilia, $, scobinate bar and cornutus. 



REVISION OF GENUS MASALIA 73 

Masalia albipuncta (Hampson) comb. n. 
(Text-figs 26, 84; PI. 8, figs 198, 199) 

Timora albipuncta Hampson, 1910 : 401. LECTOTYPE <$, Zambia (BMNH), here designated 
[examined] . 

Fore wing with areole; length (10) <J, H'2 (lectotype)-i3-8, $ (5), 120-14-4. Wing pattern 
as in PI. 8, figs 198, 199. Fore wing upper surface with ground colour pale to light yellow. 
Terminal dashes, proximal oblique-longitudinal dash (Text-fig. 26), and, when present orbicular 
and discocellular markings, reddish brown to brown. A white waved postmedial band outlined 
in reddish brown or brown may also be present. Hind wing upper surface white, distally 
suffused with pale yellow or pale yellow and light brown. 

Genitalia, o* scobinate bar and cornutus as in Text-fig. 84. ? papilla analis simple. 

Material examined. 

LECTOTYPE, here designated, [Zambia] North East Rhodesia : Upper Luangwa 
Valley, 1600-2000 ft, <£, 21.iii.1908 (S. A. Neave); paralectotype, [Zambia] North 
East Rhodesia: Upper Luangwa Valley, 1600-2000 ft, $, 16.iii.1908. 

Tanzania: Shinyanga, 1 <?, iii.1952; 1 <J, xii.1956; 4 <J, 3 ?, i.1957; 1 <J, xi.1957 
(all Croft); Shinyanga, Mwandui, 2 <$, 1 $, ii.-iii.1952 (Croft). 

Distribution. Tanzania and Zambia. 

Remarks. Apart from M . quilengesi, M. albipuncta is readily distinguished from 
other species in the genus on wing colour and pattern. M. albipuncta, which could 
be confused with M. quilengesi, can be distinguished by the presence of a proximal 
oblique-longitudinal dash on the fore wing upper surface. 




Fig. 84. M . albipuncta, $, scobinate bar and cornutus. 



74 



P. R. SEYMOUR 



Masalia quilengesi sp. n. 

(Text-fig. 85 ; PI. 8, figs 200-202) 

Fore wing with areole; length <$ (2), 11-3 (holotype)-i2-5, ? (2), n-8-i2-g. Wing pattern as 
in PI. 8, figs 200-202. Fore wing upper surface with ground colour light yellow suffused with 
pale or pastel-red. Terminal dashes, orbicular and discocellular markings brown, one, two or 
all three markings may be absent but when present from faint to well developed. A post- 
medial row of white dots, lightly differentiated, may also be present. Hind wing upper surface 
pale yellow. In a male specimen from Okahandja (PL 8, fig. 202), a distinct variant, the costa 
region is light yellow, central and anal regions greyish red and ante- and postmedial markings 
white. 

Genitalia, o* scobinate bar and cornutus as in Text-fig. 85. $ papilla analis simple. 

Material examined. 

Holotype, [Angola:] Benguella, Fort Quilenges, J, ii.1905 (Ansorge). 
Paratypes. [Angola:] Benguella, Fort Quilenges, 2 $, i. ii.1905 (Ansorge); South 
West Africa: Okahandja, 1 <J, 16. ii. 1953 (F. Gaerdes), in ZSBS, Munich. 

Distribution. Angola and South West Africa. 

Remarks. M. quilengesi is most closely allied to M. albipuncta. The anal 
proximal, oblique-longitudinal dash found in M. albipuncta and absent in 
M. quilengesi distinguishes between them. 




Fig. 85. M. quilengesi, 5*, scobinate bar and cornutus. 



REVISION OF GENUS MASALIA 



75 



Masalia terracottoides (Rothschild) comb. n. 
(Text-figs 86-88; PI. 5, figs 171-173; Map 9) 

Timora terracottoides Rothschild, 1921 : 160. Holotype £, Niger (BMNH) [examined]. 
Timora fissa Aurivillius, 1925 : 13. Holotype $, Sudan (NR, Stockholm) [examined]. Syn. n. 

Fore wing with areole present; length $ (27), 12 -8-167 (holotype 14-3), ? (20), 14-5-16-7. 
Wing pattern as in PI. 5, figs 171-173. Fore wing upper surface with variable colouring; 
ground colour from pale orange, brownish or greyish orange, light brown to brown; costal 
margin, upper central, and anal longitudinal streaks, white; postmedial dots, when present, 
brown, terminal dots white, distally tipped with brown. Hind wing upper surface white, 
immaculate or suffused with brown. 







86 




Figs 86-88. M. terracottoides genitalia. 86, $, papilla analis. 
87 and 88, q\ scobinate bar and cornutus. 



76 



P. R. SEYMOUR 



Genitalia. <J scobinate bar and cornutus as in Text-figs 87, 88. $ papilla analis modified, 
laterally flattened, with a spiculate surface (striate appearance) (Text-fig. 86). 

Material examined. 

Timora terracottoides , holotype, [Niger:] N. Damagarim, Kaleloua, <$, 8JX.1920 
(A. Buchanan); paratypes, [Niger:] Damagarim, Baban Tubki, south of Zinder, 
2 (J, 1 $, 13.ix.1920; Bande, 2 <$, 2 $, 16.ix.1920; Kaleloua, 5 <$, 9 $, 8.ix.i92o; 
Makochia, 4 <J, 3 $, 15. ix. 1920; Songo, 1 $, 17. ix. 1920 (all coll. A. Buchanan). 
Timora fissa, holotype, Sudan: [Renk,] <$, (Pr. W. Exp. Gyld), in NR, Stockholm. 

Senegal: Dakar, Aeroport, 1 <$, 9.ix.i948, in ZSBS, Munich; 1 <$, 10.ix.1956 
(C. Rungs), in MNHN, Paris; Kaolack, 2 $, 1 $, 1909 (G. Melon) ; Senegambia, 2 <^, 
x.1907 (Riggenbach) ; Gambia: i cJ (.4. Moloney). 

Distribution (Map 9). Senegal, Gambia, Niger, Angola and Sudan. 

Remarks. Except for nubila, terracottoides can be separated from all other 
species in the genus on the flattened form and spiculate surface of the papilla analis ; 
nubila differs from terracottoides in the absence of fore wing upper central and anal, 
longitudinal white streaks. 




terracottoides 



Map 9. Distribution of M. terracottoides. 






REVISION OF GENUS MASALIA 



77 



Masalia nubila (Hampson) comb. n. 
(Text-figs 89-90; PI. 8, figs 203, 204, 205) 

Timora nubila Hampson, 1903 : 108. Holotype 2, Nigeria (BMNH) [examined]. 
Timora chrysita de Joannis, 1910 : 225. LECTOTYPE $, Guinea (MNHN, Paris), here 
designated [examined]. Syn. n. 

Fore wing with areole; length <$ (4), 12 -6-13-0, $ (4), 13-4-15-0 (holotype). Wing pattern 
as in PI. 8, figs 203-205. Fore wing upper surface with ground colour light yellow to light 
orange, markings light brown to brown. Hind wing upper surface yellowish white, greyish 
orange or reddish golden brownish orange. 

Genitalia. £ scobinate bar and cornutus as in Text-fig. 90. $ papilla analis simple; surface 
spiculate (striate appearance) (Text-fig. 89). 

Material examined. 

Timora nubila, holotype [Nigeria:] River Niger between Akassa and Asaba, $ 





Figs 89-90. M. nubila genitalia. 89, $, papilla analis. 90, $, scobinate bar 

and cornutus. 



78 



P. R. SEYMOUR 



(F. D. Lugard). Timor a chrysita, LECTOTYPE, here designated. [Guinea] 
Haute Guinea; Dioudougou, $ (L.& J. de Joannis), in MNHN, Paris. 

Senegal: i <$, 6.x. 1966, in MNHN, Paris; N'Danoe, 1^, 26.viii.1951 (B. Boniface), 
in MNHN, Paris; Gambia: i $ (G. Moloney); Ghana: Northern Territories, Kete- 
Krachi, 2 <$, 1 $, {A. W. Cardinall). 

Distribution. Senegal, Guinea, Ghana and Nigeria. 

Remarks. This species is closely related to dora; the two can be distinguished 
on differences in the papilla analis, simple in nubila, modified in dora. It must, 
however, be noted that this difference is based on a single specimen of dora of which 
the abdomen has been re-attached, apparently correctly matched. 

Masalia dora Swinhoe 
(Text-figs 91, 92; PI. 8, figs 206-207) 

Masalia dora Swinhoe, 1891 : 147. LECTOTYPE <J, India (BJMNH), here designated 

[examined] . 
Timora dora (Swinhoe) Hampson, 1903 : 104. 

Fore wing with areole; length^ (1), 12-5 (lectotype), $ (1), 157. Wing pattern as in PL 8, 
figs 206-207. Fore wing upper surface with ground-colour orange to greyish orange, marked 
with brownish orange. Hind wing upper surface white. 

Genitalia. $ scobinate bar and cornutus as in Text-fig. 92. $ papilla analis modified (Text- 
fig- 9i)- 





Figs 91-92. M. dora genitalia. 91, $, papilla analis. 92, (J, scobinate bar 

and cornutus. 



REVISION OF GENUS MASALIA 79 

Material examined. 

LECTOTYPE, here designated, [India: Maharashtra,] Khandala, <J, x.1886 
(C. Swinhoe); paralectotype, [India: Maharashtra,] Khandala, 1 $ (C. Swinhoe). 

Distribution. India. 

Remarks. On pattern and colour dora appears closely allied to the African 
nubila. The striking differences between the papilla analis, modified in dora, simple 
in nubila, affords a ready means of identification. Unfortunately this difference is 
based on a single specimen of dora with a re-attached abdomen, though appearing 
correctly matched. Resemblance in other features is close but on the basis of 
limited available material, it is thought that the two names should retain their 
separate specific status. 



Masalia semifusca sp. n. 

(PI. 6, fig. 185) 

Timora dora (Swinhoe) aberration 1, Hampson, 1903 : 104. 

Timora dora ab. semifusca Warren, 1913 : 313. Holotype $, 

Timora dora ab. belgaumensis Stnmd, 1916 : 143. Holotype as above. [Objective synonym of 

semifusca'] . 

This species is known only from the female type. 

Fore wing with areole; length ? (1), 15-7 (holotype). Wing pattern (PI. 6, fig. 185). Fore 
wing upper surface colouring between light and greyish orange, almost unicolorous, though 
anal regions and scaling along the medial and radial veins is of slightly darker tone (greyish 
orange). Hind wing upper surface yellowish white, proximal area suffused with light brown. 
The specimen is slightly rubbed and probably faded. 

Genitalia. $ papilla analis simple. 

Material examined. 

Holotype, [India:] Mysore, Belgaum, $, ix.1896 (Watson), (Hampson ab. 1; ab. 
semifusca Warren holotype; ab. belgaumensis Strand holotype). 

Distribution. South-west India. 

Remarks. The relationship of semifusca to other species within the genus is 
uncertain and, with the absence of male material, its inclusion in Masalia is tentative. 
It is not thought to be conspecific with dora, as was first stated by Hampson (1903), 
nor is it thought to be closely allied, though some doubt must remain. For, although 
the papilla anali differ, simple in semifusca, modified in dora, the comparison is 
based on only two specimens of which one, the paralectotype of dora, possesses a 
re-attached abdomen. There is also the apparent absence of pattern from these 
two female specimens, but on what remains of a pattern in semifusca there is little 
to indicate close affinity. Apart from this, semifusca differs in fore wing shape and 
in possessing brown suffusion on the hind wing upper surface. 

The name semifusca was the first to be given to this unique specimen, and is 
therefore regarded here as the valid name. 



8o 



P. R. SEYMOUR 



Masalia tosta Moore 
(Text-figs 93, 94; PI. 8, figs 208, 209) 

Masalia tosta Moore, 1881 : 411. Lectotype $, India (BMNH) [examined]. 
Timora tosta (Moore) Hampson, 1903 : 115. [Lectotype designated.] 

Fore wing with areole; length <$ (1), 147, $ (5), 13-6-16-8 (lectotype). Wing pattern as in 
PI. 8, figs 208, 209. Fore wing upper surface with ground colour light, greyish, or brownish 
orange; upper central longitudinal streak, when present, yellowish white; marginal cilia, 
proximally brownish orange, distally white, the well defined line between the colours giving the 
outer margin a banded appearance. Hind wing upper surface white, yellowish white or greyish 
yellow, either immaculate or marginally suffused with light orange. 

Genitalia. <J scobinate bar and cornutus as in Text-fig. 94. $ papilla analis simple (Text- 
fig- 93)- 

Material examined. 

Lectotype, [India: Himachal Pradesh,] Dharmsala, $ {Hocking). 

India: Himachal Pradesh, Kangra, Kulu, 1 <$, 1 $ (G. Young); H.P., Sultanpur, 
3 cJ {G. Young). 

Distribution. India. 

Remarks. This species can be distinguished by its colouring, particularly the 
contrasting outer bands of the marginal cilia and, in the female, by the triangular 
shape of the papilla analis. 





Figs 93-94. M. tosta genitalia. 93, $, papilla analis. 94, <$, scobinate bar 

and cornutus. 



REVISION OF GENUS MASALIA 



81 



Masalia artaxoides (Moore) comb. n. 
(Text-fig. 95; PI. io, fig. 231) 

Pradatta artaxoides Moore, 1881 : 366. LECTOTYPE o*. India (BMNH), here designated 

[examined]. 
Timora artaxoides (Moore) Hampson, 1903 : 116. 

Fore wing with areole absent or rarely present; length $ (23), 11-9-137 (lectotype 13-2), 
$ (11), 120-170. Wing pattern as in PI. 10, fig. 231. Fore wing upper surface with ground 
colour light yellow or occasionally greyish orange; marginal cilia light yellow. Hind wing 
upper surface light yellow, yellowish grey or brownish grey; marginal cilia light yellow. 

Genitalia. <$ scobinate bar and cornutus as in Text-fig. 95. ? papilla analis simple. 

Material examined. 

LECTOTYPE, here designated, [India:] North West Himalaya, <J; paralecto- 
types, [India: Himachal Pradesh,] Dharmsala, 1 <$ {Hocking); [H.P.,], Kula, 1 $ 
{Hocking) ; [Jammu & Kashmir] Kashmir, 1 <$. 

West Pakistan: Abbottabad, 3^, 1 $, viii. (Mujtaba) ; India: Himachal Pradesh, 
Dalhousie, 1 $, v.1891 {Harford); H.P., Kangra, Kulu, 2 $, 1 $ (^4. Young), 4^ 




Fig. 95. M. artaxoides, $, scobinate bar and cornutus. 



82 P. R. SEYMOUR 

{Hocking), i $ (Swinhoe), 4$, 4 $; H.P., Kangra, Sultanpur, 4^ (G. Young) ; north- 
west India, 2 J 1 , 1 $ ; Sikkim : 1 $ (Elwes) . 

Distribution. West Pakistan, north India and Sikkim. 

Remarks. The colour and absence of pattern from the upper surface of fore and 
hind wings distinguishes artaxoides from all but four species in the genus, metaphaea, 
mittoni, modesta, and semifitsca. M. metaphaea is of darker colour and, unlike 
artaxoides, the female has a modified papilla analis. M. artaxoides can be separated 
from mittoni and modesta on differences in the scobinate bar. The bar in mittoni 
and modesta is more convex and packed closely with upward of 40 spicules (Text- 
figs 97, 98, 99) ; in artaxoides the spicules are sparsely scattered, the number rarely 
exceeding 20 (Text-fig. 95). M. artaxoides differs from seniifusca in fore wing shape 
and in hind wing marking. 

Masalia metaphaea (Hampson) comb. n. 
(Text-fig. 96; PI. 10, fig. 232) 
Timova metaphaea Hampson, 1903 : 115. Holotype $, India (BMNH) [examined]. 
The species is known only from female specimens. 

Fore wing without areole; length $ (8), 13-0-15-1 (holotype 13-4). Wing pattern as in PL 10, 
fig. 232. Fore wing upper surface with brownish orange ground colour; anal region irrorate 
with brown; marginal cilia brownish orange. Hind wing upper surface brown, marginal cilia 
brownish orange. 

Genitalia. $ papilla analis modified, terminal spines absent (Text-fig. 96). 

Material examined. 

Holotype, [India: Himachal Pradesh,] N. W. Himalaya, Mandi, 3-5000 ft, $, 
1883 {G. Young). 

West Pakistan: Baluchistan, 1 $; India: Himachal Pradesh, Mandi, 3-5000 ft, 
3 $, 1883 (G. Young); Sikkim, 3 $ {Elwes). 

Distribution. West Pakistan, north-east and north-west India, Sikkim. 

Remarks. Upper surface fore and hind wing colouring and virtual absence of 
pattern (fore wing anal region irrorate with brown) distinguishes metaphaea from all 
other species in the genus. 

Masalia mittoni (Pinhey) comb. n. 
(Text-figs 97, 98 ; PI. 10, fig. 233) 

Timora mittoni Pinhey, 1956 : 13. LECTOTYPE <$, Tanzania (BMNH), here designated 
[examined] . 

Fore wing with areole; length 3 (18), 12-1-13-9 (lectotype 13-5), $ (14), I37-I5-5- Wing 
pattern as in PI. 10, fig. 233. Fore wing upper surface with ground colour from pale to light 
orange, light to moderately irrorate with brown; extent of irrorate area variable. Hind wing 
upper surface light brown to brown. 

Genitalia. 6* scobinate bar and cornutus as in Text-figs 97, 98. $ papilla analis simple. 






REVISION OF GENUS MASALIA 



83 



Material examined. 

LECTOTYPE, here designated, [Tanzania] Tanganyika: Iringa, <$, iii.1950 
(N. Milton); paralectotypes, [Tanzania] Tanganyika: Iringa, 11 <$, 14 $, iii.1950 
{N. Mitton). 

Tanzania: Iringa, 6^, iii.1950 (N. Mitton). 

Distribution. Tanzania. 

Remarks. The fore and hind wing colouring of the upper surface, and absence 
of a definite pattern, distinguish mittoni from all but two other species in the genus, 
metaphaea and artaxoides. M. metaphaea is of darker colouring and, unlike mittoni, 
the female has a modified papilla analis. M. artaxoides and mittoni can be separated 
on differences in the scobinate bar. The bar in mittoni is more convex and closely 
packed with upward of 40 spicules (Text-figs 97, 98) ; in artaxoides the spicules are 
sparsely scattered and rarely exceed 20 in number (Text-fig. 95). 




'"vm 



If ail 

1 1 ' 



' ' ' 


^<0- 


ILm^ 




. \ 1 





96 



I 



Fig. 96. M . metaphaea, $, papilla analis. 



8 4 



P. R. SEYMOUR 



Masalia modesta (Moore) comb. n. 
(Text-fig. 99; PI. 10, fig. 234) 

Pradatta modesta Moore, 1881 : 366. LECTOTYPE ? India (BMNH) here designated 

[examined] . 
Timor a modesta (Moore) Hampson, 1903 : 116. 
Curubasa calamaria Moore, 1881 : 367. LECTOTYPE <J, India (BMNH), here designated 

[examined]. [Synonymized by Hampson 1903 : 116.] 

Fore wing with areole; length o* (3). 12-0-12-3, ? ( l )> II- ° (lectotype). Wing pattern as in 
PI. 10, fig. 234. Fore wing upper surface pale yellow. Hind wing upper surface pale yellow. 
Genitalia, o* scobinate bar and cornutus as in Text-fig. 99. $ papilla analis simple. 

Material examined. 

Pradatta modesta, LECTOTYPE, here designated, India: [Uttar Pradesh, Main- 
puri] Manpuri, $; Curubasa calamaria, LECTOTYPE, here designated, [India: 
Maharashtra,] Bombay, <$. 

India: Madhya Pradesh, Jubbulpore, 2 $. 

Distribution. India. 

Remarks. The four specimens in the collection of the British Museum (Natural 
History) appear faded. In one specimen the fore wing upper surface has traces of 
pale red on the costal and anal regions. 

The colour and virtual absence of pattern from the upper surface of the fore and 
hind wings separate modesta from all but one other species in the genus, artaxoides. 
M. modesta and artaxoides can be separated on differences in the scobinate bar. 
The bar in modesta is more convex and closely packed with upward of 40 spicules 
(Text-fig. 99) ; in artaxoides the spicules are sparsely scattered and rarely exceed 20 
in number (Text-fig. 95). 





\j- -^ 

'1 '■■ ,■■■ > \ \ 



97 




Figs 97-98. M. mittoni, <$, scobinate bar and cornutus. 



REVISION OF GENUS MASALIA 



85 



Masalia bimaculata (Moore) comb. n. 
(Text-figs 100-109; PI- 9< fig s 210-221; Map 10) 
Pradatta bimaculata Moore, 1888 : 411. 

M. bimaculata is a variable species, in size, extent of pattern differentiation and 
colour. A combination of two characters, (1) the presence of a brown discocellular 
spot, occasionally faint but rarely indistinguishable against the surrounding colour, 
and (2) the modified form of the papilla analis devoid of terminal spines, separates 
M . bimaculata from other species in Masalia. 

Four subspecies are recognized, the structural differences being as follows : 

Areole 

present 

absent 

present or absent 

absent 

Though variation in colour and marking occurs between the four, pattern differ- 
ences are not marked and are further obscured by variation within, and slight 
overlap between, them. M. b. nigrifasciata, linking with M. b. bimaculata and 
M. b. cornia on presence and absence, respectively, of an areole is, on colour and mark- 
ing, the most readily distinguished. These differences are set out below. 

M. b. nigrifasciata. Costal and anal regions typically dull red, central region yellow- 
ish white to light yellow; dark brown lower central longitudinal streak well 
developed. 



M. b. bimaculata 
M. b. cornia 
M. b. nigrifasciata 
M. b. pluritelifora 



Cornutus 


Distribution 


present 


Indian 


present 


African 


present 


African 


absent 


African 




Fig. 99. M. modesta, <$, scobinate bar and cornutus. 



86 



P. R. SEYMOUR 



M. b. bimaculata. Costal, central and anal regions white to pale yellow; brown 

lower central longitudinal streak absent or present. 
M. b. cornia. Costal, central and anal region from pale yellow to reddish grey; 

dark brown lower central longitudinal streak present or absent. Or, costal 

and anal regions reddish white, with a poorly developed brown lower central 

longitudinal streak. 





Figs 100-103. M. bimaculata subspecies, scobinate bar and cornutus of male genitalia. 
100, bimaculata bimaculata. 101, bimaculata nigrifasciata. 102, bimaculata pluritelif or a. 
103, bimaculata cornia. 



REVISION OF GENUS MASALIA 87 

M. b. pluritelifora. Costal, central and anal regions greyish orange to reddish grey 
irrorate with brown; a white upper central longitudinal streak usually present; 
brown lower central longitudinal streak poorly developed or absent. 



Masalia bimaculata bimaculata (Moore) 
(Text-figs 100, 104; PI. 9, figs 210, 211; Map 10) 

Pradatta bimaculata Moore, 1888 : 411. Holotype o", India (BMNH) [examined]. 

Timora bimaculata (Moore) Hampson, 1903 : 109. 

Pradatta pallescens Hampson, 1891 : 70. LECTOTYPE <J, India (BMNH), here designated 
[examined]. [Synonymized by Hampson, 1903 : 109.] 

Pradatta pulverulenta Hampson, 1891 : 71. Holotype <J, India (BMNH) [examined]. [Syn- 
onymized by Hampson (stated to be an aberration), 1903 : 109.] 

Fore wing with areole; length o* ("). 104 (hoIotype)-i4-6, $ (5), 132-155. Wing pattern 
as in PI. 9, figs 210, 211. Fore wing upper surface with ground colour yellowish white to pale 
yellow, immaculate or finely irrorate with brown. Lower central longitudinal streak (when 
present) and discocellular spot, brown. Hind wing upper surface white to yellowish white. 

Genitalia. <£ scobinate bar and cornutus as in Text-fig. 100. ? papilla analis modified, 
terminal spines absent, dorso-lateral surface sericate (Text-fig. 104). 

Material examined. 

Pradatta bimaculata, holotype, [India: Himachal Pradesh,] Dharmsala, $ 
{J.H.Hocking). Pradatta pallescens, LECTOTYPE, here designated, [India: Madras,] 
Nilgiris, $, {Hampson); paralectotypes, [India: Madras,] Nilgiris, 3 <$ {Hampson). 
Pradatta pulverulenta, holotype, [India: Madras,] Nilgiris, <J {Hampson). 

India: Himachal Pradesh, Kangra Valley, 4500 ft, 1 <J, ix.1899 {Dudgeon); H.P., 
Sultanpur, Kulu, 1 <$; H.P., Sultanpur, Kulu, 1 $, 1889 {G. Young); Kerala, Peer- 
made, 1 $ {Imray); K., Travancore Place, 1^,1$; Madras, Nilgiris, 1 <$, 1 $; M., 
Nilgiris, 1 £ {Hampson) ; Madhya Pradesh, Mhow, 1 $. 

Distribution (Map 10). India. 

Remarks. The nominate subspecies and M. b. nigrifasciata are distinguishable 
on differences in colouring of the costal and anal regions, and the presence of the 
areole in M. b. bimaculata distinguishes it from M . b. cornia and M. b. pluritelifora, 
in which areoles are absent. 



Masalia bimaculata nigrifasciata (Hampson) comb, et stat. n. 

(Text-figs 101, 108, 109; PI. 9, figs 216-219; Map 10) 

Timora nigrifasciata Hampson, 1903 : no. LECTOTYPE q\ Kenya (BMNH), here designated 

[examined]. 
Timora bimaculata var. unifasciata Gaede, 1915 : 39. Holotype q\ Tanzania (MNHU, Berlin) 

[examined]. [Synonymized with Timora pulverulenta Hampson by Gaede, 1935 : 106.] 

Syn. n. 



88 P. R. SEYMOUR 

Fore wing with or without areole ; length^ (83), 13-0-16-5 (lectotype 15-8), $ (42), 14-9-18-1. 
Wing pattern as in PI. 9, figs 216-219. Fore wing upper surface with costal and anal regions 
pale to dull red ; central region yellowish white to light yellow occasionally with pale to dull red 
extending in from the costal and anal regions. In a number of specimens the regions are finely 
irrorate with dark brown; lower central longitudinal streak and discocellular spot also dark 
brown. Hind wing upper surface yellowish white to pale yellow. 

Genitalia. 5* scobinate bar and cornutus as in Text-fig. 101. $ papilla analis modified, 
terminal spines absent, dorso-lateral surface sericate (Text-figs 108, 109). 



Material examined. 

Timora nigrifasciata, LECTOTYPE, here designated, [Kenya] B.E. Africa: Eb 
Urru, <$, 16.iv.1900 (C. S. Betton); paralectotypes, [Kenya] B.E. Africa: Eb Urru, 
1 <J ( 21.iii.1900; 1 $, 22.iii.1900; 1 $, 23.iii.1900; 1 $, i.iv.1900; 1 (J, 20.iv.1900, 1 $, 
30. v. 1900 (all C. S. Betton). Timora bimaculata var. unifasciata, holotype, [Tan- 
zania:] Mkalama, <J, 4.1V.1905 (Marwitz), in MNHU, Berlin; paratype, [Tanzania:] 
Mkalama, $, 9.11.1905 (Marwitz), in MNHU, Berlin. 

Uganda: W. Ankole, 4500-5000 ft, 1 $, n.x.1911 (S. A. Neave); Mbarara, 1 6* 
(R. E. McConnell) ; S.E. Ruwenzori, 3500 ft, 1 <$, 16.iv.1906 (G. Legge & A. F. R. 
Wollaston); Kenya: Eb Urru, 2 $, 22.iii.1900; 1 $, 23.iii.1900; 1 <$, 25.iii.1900; 1 <$, 
26.iii.1900; 1 $, 31.iii.1900; 1 <$, 19.iv.1900 (all C. S. Betton); Mt. Elgon, 1 $, v.1932 
(T. H. E. Jackson) ; Hoey's Bridge, 2 $ [Pitman) ; Kavirondo, Suna, 1 <$, 1 $, xi.1930; 
1 $, i.1931; 1 (J, iii.1931; 1 (J, 2 $, iv.1931; 4 (J, xi.1931; 10 cJ, xii.1931; 7(J, 2 $, 
i.1932; 1 <J, ii.1932; 3 & 2 $, iii.1932; 9 (J, 4 $, iv.1932; 2 (J, 2 $, v.1932 (all 
W. Feather) ; 2 <$, 7 $, iv.1932 (w<m Someren) ; Kitale, 1 (J, 24.iv.1927; 1 (J, 17.lv. 1931; 
1 (J, (all G. W. Jeffery); Lumbwa, 1 ^, 10.iv.1923 (G. W. Jeffery); Nairobi, 1 <$, 
i.vi.1905 (F. J. Jackson) ; 2 $, 21.iv.1916 (W 7 . yl. Lamborn) ; 5600 ft, 3 ^, 1 $ (PF. 2V. 
wan Someren) ; 1 <j>, iv.1927 (D. M. Hopkins) ; Nakutu, 1^,1$, 8.V.1911 ; 1 <$ (H. A. 
Bodeker) ; 2 ^, iv.1940 ; 2 (J, iv.1941 ; 1 $, 6.VL1943 ; 1 <$, 17.viii.1944 (all A . Townsend) ; 
Nandi, Moboroni, 1 $, vii.1903 (F. J. Jackson); Tanzania: Arusha District, 1 $ 
(M. S. Moore) ; Arusha District, Ngorongoro Crater, 5900 ft, 1 <$, 2 $, iii.1921 (T. A. 
Barns) ; District of Great Craters, 1 $, ii— iii.1921 (T. A. Barns) ; Kigoma, 1 $, iv.1961 
(Goodall); W. Kilimanjaro, 4000-5000 ft, 1^,1$, ii.iii.1937; 1 $, iv.-v.1937 
(B. Cooper); Musoma, Banagi Hill, 1 <$ (M. S. Moore); Nachingwe, 2 $, iv.1961 
(W. Bigger) ; Ethiopia : 1 $. Specimens with incomplete data 4 $, 2 $. 

Distribution (Map 10). Uganda, Kenya, Tanzania and Ethiopia. 

Remarks. M. b. nigrifasciata is distinguishable from the nominate subspecies 
and from M. b. pluritelifora on the colour difference of the costal and anal regions. 
Difference in colour and marking also serves to distinguish most specimens of 
M. b. cornia from M. b. nigrifasciata. The range of variation in these two subspecies 
however, is such that a continuous series can be traced from one to the other. In 
M. b. nigrifasciata there is a shift of tone from dull red toward greyish orange, 
accompanied by dilution of dark brown from the lower central longitudinal streak 
(PI. 9, figs 216, 217). The known intergrade variants were taken from localities on 
the common borders between the two subspecies. 



REVISION OF GENUS MASALIA 



89 




•' \ : : l ■ ■ 









104 



105 



106 





V.M: 



109 

/ 



Figs 104-109. M. bimaculata subspecies, papilla analis of female genitalia. 104, bimacu- 
lata bimaculata. 105, bimaculata cornia. 106, bimaculata cornia. 107, bimaculata 
pluritelifora. 108, bimaculata nigrifasciata. 109, bimaculata nigrifasciata. 



go P. R. SEYMOUR 

Masalia bimaculata cornia subsp. n. 
(Text-figs 103, 105, 106; PI. 9, figs 212-215; Map 10) 

Fore wing without areole; length £ (10), 10-5-14-5 (holotype), $ (2), 12-3-15-4. Wing 
pattern variable (PI. 9, figs 212-215). Fore wing upper surface with costal, central and anal 
regions yellowish white to pale or greyish orange, finely, moderately or densely irrorate with 
brown. Central region uniform or of different colour to costal and anal regions. Discocellular 
spot, and lower central longitudinal streak when present, brown. A white upper central 
longitudinal streak may also be present. Hind wing upper surface yellowish white. 

Genitalia, o* scobinate bar and cornutus as in Text-fig. 103. $ papilla analis modified, 
terminal spines absent, dorso-lateral surface seriate (Text-figs 105, 106). 

Material examined. 

Holotype, [Congo (Kinshasa):] Luvua River (east Bank), 85 miles North of 
Lake Mweru, 3000 ft, <$, iv.1922 (end of wet season) (T. A. Barns). 

Paratypes. [Angola:] Capelongo, 1 <$, 20.xii.1912 (Mission Rohan-Chabot), in 
MNHN, Paris; [Congo (Kinshasa):] Luvua River, 85 miles north of Lake Mweru, 
3000 ft, 1 $, 1 $, ix.1922 (T. A. Barns) ; [Sudan:] Darfur Province, Kulme, 1 $, 1921 
(H. Lynes) ; Gondokoro, White Nile, 5 <£, 1 $ (W. E. Reymes-Cole) ; Southern Bahr-el- 
Ghazal, Tambura, 1 <J. 

Distribution (Map 10). Congo (Kinshasa) and Sudan. 

Remarks. The presence of a cornutus in this subspecies readily distinguishes it 
from M. b. pluritelifora, whilst the absence of an areole distinguishes it from M . b. 
bimaculata. M. b. cornia and M. b. nigrifasciata differ in fore wing colour and pattern, 
but both are variable and intergrade variants do occur. 

The name cornia was a manuscript name of Miss A. E. Prout and the type selected 
by her is designated here as holotype. 



Masalia bimaculata pluritelifora (Berio) comb, et stat. n. 
(Text-figs 102, 107; PI. 9, figs 220, 221; Map 10) 

Timora pluritelifora Berio, 1966 : no. Holotype $, Senegal (MNHN, Paris) [examined]. 
Timora rosastrigata Berio, 1966 : in. Holotype $, Senegal (MNHN, Paris) [examined]. 
Syn. n. 

Fore wing without areole; length 5* (8), n-o-n-8, $ (8), 12-2-14-1 (holotype 13-2). Wing 
pattern as in PL 9, fig. 220, 221. Fore wing upper surface with ground colour greyish orange, 
greyish red, pale red or reddish grey, fine to densely (as in type) irrorate with brown. When 
present, upper central longitudinal streak white; discocellular spot brown, faint (occasionally 
indistinguishable against the irrorate ground) to well developed. Hind wing upper surface 
yellowish white. 

Genitalia. $ cornutus absent; scobinate bar and proximal end of vesica as in Text-fig. 102. 
$ papilla analis modified, terminal spines absent, dorso-lateral surface sericate (Text-fig. 107). 



REVISION OF GENUS MASALIA 



9i 



Material examined. 

Timora pluritelifora, holotype, Senegal: Badi, Pare National Niokola Koba, $, 
15.vii.-25.ix.1955 (Instit. franc. d'Afrique noire, Dakar), in MNHN, Paris. Timora 
rosastrigata, holotype, Senegal: Badi, Pare National Niokola Koba, 9. 3-i XI 958 
(Inst, franc. d'Afrique noire, Dakar), in MNHN, Paris. 

1 Senegal: Dakar, 3 <J, 1 $, 10-12.ix.1956 (C. Rungs), in MNHN, Paris; Kaolack, 
1 $ 1909 (G. Melon); Sedhiou, 1 <J, 1 $, 1917 ( H - CasteU); Ghana: Northern Terri- 
tories, Kete-Krachi, 2 <$, 4 $ {A. W. Cardinall) ; Nigeria: Ogruga [? Ogrugru], River 
Niger, 2 <J, 1 $. 

Distribution (Map 10). Senegal, Ghana, Nigeria. 

Remarks. The absence of a cornutus in the male of M. b. pluritelifora distin- 
guishes it from the three other subspecies. 




• ..,.., m bimaculata cornia 

bimaculata bimaculata 

A bimaculata nigrifasciata ♦ bimaculata pluritelifora 

Map 10. Distribution of subspecies of M. bimaculata. 



92 



P. R. SEYMOUR 



Masalia flavocarnea (Hampson) comb. n. 
(Text -fig. no; PI. 9, fig. 222) 
Timora flavocarnea Hampson, 1903 : 115. Holotype ?, Ethiopia (BMNH) [examined]. 

Fore wing without areole; length $ (1), 16-8 (holotype). Wing pattern as in PI. 9, fig. 222. 
Fore wing upper surface with ground colour pale to greyish orange, discocellular spot brown. 
Hind wing upper surface pale yellow irrorate with light brown. 

Genitalia. $ papilla analis modified; dorso-lateral surface sericate and with terminal spines 
(Text-fig. no). 

Material examined. 

Holotype, [Ethiopia] Abyssinia, $. 

Distribution. Ethiopia. 

Remarks. Within the genus, M. flavocarnea can be separated from all but 
M. bimaculata on fore wing colour and marking. M. flavocarnea and M. bimaculata 
are separable on the presence and absence respectively of papilla anali terminal 
spines. 







< / Nm ^ no 



Fig. no. M. flavocarnea, $, papilla analis. 



REVISION OF GENUS MASALIA 



93 



Masalia albida (Hampson) comb. n. 
(Text-figs 20, in; PI. 10, figs 223-225) 

Timora albida Hampson, 1905 : 450. LECTOTYPE q\ Algeria (BMNH), here designated 
[examined]. [Synonymized with Argyrospila striata Staudinger, 1897 : 265, by Warren, 
191 1 : 248; recalled from synonymy by Draudt, 1935 : 197.] 

Lecerfia chitinipyga Dumont, 1920 : 102. LECTOTYPE q", Algeria (MNHN, Paris), here 
designated [examined]. [Synonymized by Draudt, 1935 : 197.] 

Antenna with flagellar segments sexually dimorphic. Proboscis quotient: 6. Fore wing 
with areole; length q* (51). i3"2-i5-8 (lectotype 14-3), ? (40), 13-9-16-4. Wing pattern as in 
PI. 10, figs 223-225. Fore wing upper surface with ground colour greyish yellow to greyish 
orange. Upper central and anal longitudinal streaks, and costal and anal margins, white. In 
addition a variable number of white streaks may be present; veins between which streaks occur 
are: R4-R5, R5-M1, M2-M3, Mz-Cui & and CMia-C^ib- Hind wing upper surface white to 
yellowish white, either immaculate or central veins greyish orange. Eighth abdominal tergum 
with posterior margin centrally incurved; incurved region ridged (Text-fig. 20). 

Genitalia, o* scobinate bar and cornutus as in Text-fig. ill. $ papilla analis simple. 

Material examined. 

Timora albida, LECTOTYPE, here designated, Algeria: Hammam-es-Salahin, 
o*, 9.1V.1904; paralectotype, Algeria: Hammam-es-Salahin, $, 16. iv. 1904. Lecerfia 
chitinipyga, LECTOTYPE, here designated, [Algeria] Algine: El Golea, <$, in 
MNHN, Paris. 

Algeria: east of Guerrara, 1 <$, 13.iv.1914; El Alia, between Touggourt and 
Guerrara, 1 $, 4 $, 12.iv.1914; Hassi Dinar, south of Touggourt, 1 $, 11.iv.1914; 
Hassi Sidi Mahmud, between El Arich and Oued Nga, 1 <$, 4.^.1914; Mzab Country, 
Oued Nga, 1 $, 16-30.iv.1914; South Oran, Ain Sefra, 31 £, 15 $; 3-9. v. 191 3;2 (J, 



1 




Fig. hi. M. albida, $, scobinate bar and cornutus. 



94 



P. R. SEYMOUR 



i $, 13.V.1915 (V. Faroult); Touggourt, 1 <$, 12.iv.1914; Saudi Arabia: El Riad, 
3 cJ, 5 ?, xi.1958 (E. Diehl), in ZSBS, Munich; El Ryadh, 3 <J, 6 $, 18.iii.1958 
(£. Zta«), in ZSBS, Munich; Jidda, 1 (J, 13.iii.1929 (#. Si/. 5. P/w'%) ; Tihama of 
Asir, 1 $, 27.iii.1948; Iran: Belutschistan, Jranshar, 800 m, 3 $, 3 $, 1-10.iii.1954 
(Richter & Schauffele) ; 1 $, 28-31. iii. 1954 (Richter & Schauffele), in ZSBS, Munich. 

Distribution. Algeria, Arabia and Iran. 

Remarks. The long tongue together with the ridged incurved region of the 8th 
abdominal tergum readily distinguishes albida from other species in this genus. 



Masalia per striata (Hampson) comb. n. 
(Text-figs 10, 112, 113; PI. 10, figs 226-230; Map 11) 

Raghuva perstriata Hampson, 1903 : 32. 

Antenna with flagellar segments sexually dimorphic. Proboscis quotient: 5. Fore wing 
with areole. In the male, the costa dilates at a point a little beyond the centre to form a node. 
Into this node runs an elongate ridge, which arises from a position anterior to the areole, between 
R1-R4 (Text-fig. 10). The node and ridge are not differentiated in the female. 

Genitalia, o* scobinate bar and cornutus as in Text-figs 112, 113. $ papilla analis simple. 

Remarks. The long proboscis together with male characters, the node and ridge 
in the fore wing and the few but well developed scobinate bar spicules, distinguish 
M. perstriata from other species in the genus. 

Three subspecies are recognized, the nominate subspecies from India, M. p. 
fuscostriata from Arabia and Iran, and M. p. zernytamsia from east Africa. The 
nominate subspecies and M. p. zernytamsia are marked on the fore wing with a row 
of terminal dots, which are absent in M. p. fuscostriata. No morphological differ- 
ences have been found between M. p. perstriata and M. p. zernytamsia. 





Figs 112-113. M. perstriata subspecies, scobinate bar and cornutus of male genitalia. 
112, perstriata perstriata. 113, perstriata fuscostriata . 



REVISION OF GENUS MAS A LI A 95 

Masalia perstriata perstriata (Hampson) 

(Text-fig. 112; PI. 10, fig. 229, Map 11) 

Raghnv a perstriata Hampson, 1903 : 32. Holotype <$, India (BMXH) [examined]. 

This subspecies is known only from male specimens. 

Fore wing length, $ (2), 13-8-14-0 (holotype). Wing pattern as in PI. 10, fig. 229. Fore 
wing upper surface with ground colour yellowish white to pale yellow; postmedial and terminal 
dots and the rather faintly marked lower central, longitudinal streak, brown. Hind wing 
upper surface white to yellowish white. 

Material examined. 

Holotype, India: [Himachal Pradesh.], N. W. Himalayas, Fort Kangra, <J, 
17.vii.1884 {Moore). 

India (southern): Gooty, 1 <$ (W. H. Campbell). 

Distribution (Map n). India. 

Remarks. The presence in M. p. perstriata of brown terminal dots distinguishes 
it from M. p. fuscostriata, in which these dots are absent. Morphological differences 
have not been found between M. p. perstriata and M. p. zernytamsia. These two 
subspecies are geographically separated by M. p. fuscostriata (Map 11). 







perstriata perstriata 
perstriata fuscostriata 
perstriata zernytamsia 



Map 11. Distribution of subspecies of M. perstriata. 



96 P. R. SEYMOUR 

Masalia perstriata fuscostriata (Brandt) comb, et stat. n. 

(Text-fig. 113; PI. 10, figs 226-228; Map 11) 

Timora fuscostriata Brandt, 1914 : 854. LECTOTYPE <$, Iran (NR, Stockholm), here desig- 
nated [examined]. 

Fore wing, length <J (3), 14-3— 15-4 (lectotype), $ (3), 15-2-17-0. Wing pattern as in PI. 10, 
figs 226-228. Fore wing upper surface with ground colour white to yellowish white; costal and 
anal regions fine to moderately irrorate with brown; lower central longitudinal streak either 
straight or distad-splayed and postmedial dots, when present, brown. Hind wing upper 
surface white to yellowish white. 

Material examined. 

LECTOTYPE, here designated, Iran: Baloutchistan, Bender Tchahbar, <$, 
27.ii.-3.iii.1938 (Brandt), in NR, Stockholm; paralectotypes, Iran: Baloutchistan, 
Bender Tchahbar, 1^,1$, 27.ii.-3.iii.1g38 (Brandt), in NR, Stockholm; 1 <$, 1 9, 
16 and 20. ii. 1938 (Brandt), in ZSBS, Munich. 

Saudi Arabia: Rada, 1 $, 4.1X.1962 (G. Popov). 

Distribution (Map 11). Saudi Arabia and Iran. 

Remarks. The absence of fore wing terminal dots in M. p. fuscostriata distin- 
guishes it from the two other subspecies, M. p. perstriata and M. p. zernytamsia, in 
which brown terminal dots are present. 



Masalia perstriata zernytamsia (Berio) comb, et stat. n. 
(PI. 10, fig. 230; Map 11) 

Timora zernytamsia Berio, 1939 : 60. Holotype $, Somali Republic: Belet, 15.vii.1934 
(Patrizi) (MCSN, Genoa). 

Fore wing, length $ (12), 11-8-14-7, $ ( J 4). i 4' 0_i 5'5- Wing pattern as in PL 10, fig. 230. 
Fore wing upper surface with ground colour yellowish white to pale yellow, anal region occa- 
sionally irrorate with brown ; postmedial and terminal dots and lower central longitudinal streak 
brown. Hind wing upper surface white to yellowish white. 

Material examined. 

Ethiopia: El Dire, Sagan-Omo, 1 $, in MCSN, Genoa; Dire Daoua, 10 <$, 13 $, 
xii.1934, or iv.-ix.1935 (H. Uhlenhuth); Kenya: Isiolo, 2$, (H. Copley). 

Distribution (Map 11). Somali Republic (type-locality), Ethiopia and Kenya. 

Remarks. Morphological differences between M. p. zernytamsia and M. p. 
perstriata have not been found. They occur in different regions, M. p. zernytamsia 
in Africa, M . p. perstriata in India. Lying between them geographically is the third 
subspecies, M. p. fuscostriata (Map 11). M. p. zernytamsia and M. p. fuscostriata 
are separable on the presence and absence, respectively, of a row of fore wing 
terminal dots. 



REVISION OF GENUS MASALIA 97 

REFERENCES 

Aurivillius, C. 1925. Zoological Results of the Swedish Expedition to Central Africa 1921. 

(Lepidoptera). Ark. Zool. 17A (32) : 1-20. 
Berio, E. 1935. Spedizione zoologica del Marchese Saverio Patrizi nel Basso Giuba e 

nell' Oltregiuba. 1934- Nuove specie di Eteroceri. Annali Mus. civ. Stor. nat. Giacomo 

Doria 58 : 56-65, 7 figs. 
1941. Contributi alio studio dei Lepidotteri Eteroceri dell' Eritrea. VII. Euchromiidae, 

Arctiidae, Agaristidae, Lymantriidae, Lasiocampidae, Noctuidae raccolti dal Sig. G. 

Vaccaro nel 1938. Annali Mus. civ. Stor. nat. Giacomo Doria 61 : 176-190. 

IQ 53- Contributo alia Conoscenza di Noctuidae poco note, diagnosi di nuove specie e 

note critiche. Doriana 1 (34) : 1-6, 13 figs. 

1962. Diagnosi di Alcune Specie di Noctuidae Africane. Boll. Soc. ent. Hal. 92 : 122-126, 

8 figs. 

1966. Descrizione di Nuove Noctuidae Africane e Note Sinonimiche. Annali Mtts. civ. 

Stor. nat. Giacomo Doria 76 : 1 10-136. 

Bethune-Baker, G. T. 191 1. Descriptions of new species of Lepidoptera from Tropical 

Africa. Ann. Mag. nat. Hist. (8) 8 : 506-542. 
Boursin, C. i960. Nouvelles "Trifinae" d'Afghanistan de l'Expedition Klapperich (3 mc 

note) (Lep. Noctuidae) (Diagnoses preliminaires). Bull. mens. Soc. linn. Lyon 29 (5) : 

136-152. 
Brandt, W. 1941. Beitrag zur Lepidopteren-Fauna von Iran (3). Mitt, munch, ent. Ges. 

31 : 835-863. 
Butler, A. G. 1886. Descriptions of 21 new genera and 103 new species of Lepidoptera- 

Heterocera from the Australian Region. Trans, ent. Soc. Lond. 19 : 381-441, 2 pis. 
de Joannis, J. 1910. Description de trois nouvelles especes de Timora [Lep. Noctuidae] 

provenant de la Haute-Guinee francaise. Bull. Soc. ent. Fr. 1910 : 223-226. 

1913. Materiali per lo Studio della Fauna Eritrea raccolti nel 1901-03 dal Dott. A. 

Andreini. Lepidopteres. Heterocera. Bull. Soc. ent. ital. 44 (191 2) : 122-147, 4 figs. 

Distant, W. L. 1902. Descriptions of new species of Heterocera from the Transvaal. 

Entomologist 35 : 212-214. 
Draudt, M. 1935. In Seitz, Macrolepidoptera of the World. Suppl. 3 : 197, pi. 22. 

Stuttgart. 
Druce, H. 1887. Descriptions of some new species of Lepidoptera Heterocera, mostly from 

Tropical Africa. Proc. zool. Soc. Lond. 1887 : 668-686, pi. 60. 
1889. Noctuidae. Biologia cent.-am. 1 : 257-423, pi. 26, fig. 2-pl. 34, fig. 3. 

1903. Descriptions of some new species of Lepidoptera, chiefly from South America. 

Ann. Mag. nat. Hist. (7) 11 : 196-203. 

Dumont, C. 1920. Contribution a l'etude des Lepidopteres du Sahara algerien. Description 

d'une espece nouvelle de Trifinae. [Lep. Noctuidae]. Bull. Soc. ent. Fr. 1920 : 102-104, 

1 fig. 
Gaede, M. 1915. Neue und wenig bekannte afrikanische Timora-Arten (Fam. Agrotinae). 

Int. ent. Z. 9 : 39-40. 

■ 1935. In Seitz, Macrolepidoptera of the World 15 : 105-108, pi. 11, 17 figs. Stuttgart. 

Grunberg, K. 1910. Lepidoptera. In Schultze, L. G. Zoologische und anthropologische 

Ergebnisse einer Forschungsreise in westlichen und zentralen Sudafrika, 1903-05. Denkschr. 

med.-naturw. Ges. Jena 16 : 91-146, pi. 3, 4 figs. 
Hampson, G. F. 1891. Illustrations of typical specimens of Lepidoptera Heterocera in the 

Collection of the British Museum 8 : 1-144, pis 139-146. London. 

1902. The Moths of South Africa 2. Ann. S. Afr. Mus. 2 : 255-446. 

1903. Catalogue of the Lepidoptera Phalaenae in the British Museum 4 : 666 pp., 125 figs. 

London. 
• 1905. Descriptions of new genera and species of Syntomidae, Arctiadae, Agaristidae, and 

Noctuidae. Ann. Mag. nat. Hist. (7) 15 : 425-453. 



98 



P. R. SEYMOUR 



Hampson, G. F. 1907. Descriptions of new genera and species of Syntomidae, Arctiadae, 
Agaristidae, and Noctuidae. Ann. Mag. nat. Hist. (7) 19 : 221-257. 

iqio. Zoological Collections from Northern Rhodesia and adjacent Territories: Lepi- 

doptera Phalaenae. Proc. zool. Soc. Lond. 1910 : 388-510, pis 36-41. 

Hardwick, D. F. 1965. The Corn Earworm Complex. Mem. ent. Soc. Can. 40. 
Methuen. Handbook of Colour. Second Edition, 1967. pp. 243, 30 col. pis. London. 
Moore, F. 1881. Descriptions of new genera and species of Asiatic Nocturnal Lepidoptera. 
Proc. zool. Soc. Lond. 1881 : 326-380, pis 37-38. 

1888. Descriptions of new genera and species of Lepidoptera Heterocera collected by 

Rev. J. H. Hocking, chiefly in the Kangra District, N.W. Himalaya. Proc. zool. Soc. 
Lond. 1888 : 390-412. 

Pinhey, E. C. G. 1955. Some new species of Lepidoptera from Eastern Africa. Occ. Pap. 

Coryndon meml Mus. 4 : 10-16, pi. 1. 
Prout, A. E. 192 1. New Lepidoptera collected by Mr T. A. Barns. III. New Noctuidae. 

Bull. Hill Mus. Witley 1 : 1 19-138, pi. 17. 
Rothschild, W. 192 i. On the Lepidoptera collected by Captain A. Buchanan in Northern 

Nigeria and the Southern Sahara in 1919-1920. Novit. zool. 28 : 142-170. 
Staudinger, O. 1897. Vier neue Heteroceren aus Algerien und Tunesien. Dt. ent. Z. Iris 

10 : 265-270, pi. 4, fig. 4. 
Strand, E. 1916. Neue Aberrationen der Noctuiden Subfamilien Agrotinae und Cuculiinae. 

Arch. Naturgesch. 81 (A12) : 142-149. 
Swinhoe, C. 1885. On the Lepidoptera of Bombay and the Deccan. Part III, Heterocera. 

Proc. zool. Soc. Lond. 1885 : 447-476, pis 37-38. 

1891. New species of moths from Southern India. Trans, ent. Soc. Lond. 24 : 133-154, 

pi. 8. 

Viette, P. 1957. Descriptions preliminaires de nouvelles especes de Noctuelles de Madagascar 
I. [Lep. Noctuidae]. Bull. Soc. ent. Fr. 62 : 270-279. 

1958. Descriptions preliminaires de nouvelles especes de Noctuelles de Madagascar II. 

[Lep. Noctuidae]. Bull. Soc. ent. Fr. 63 : 146-152. 

Walker, F. 1856. List of the specimens of Lepidopterous insects in the Collection of the British 

Museum 9 : 1-252. London. 
1865. List of the specimens of Lepidopterous insects in the Collection of the British Museum. 

33 : 707-1120. London. 
1866. List of the specimens of Lepidopterous insects in the Collection of the British Museum. 

35 : 1535-1984. London. 
Wallengren, H. D. J. 1856. Anteckningar i Zoologien. Lund. 
Warren, W. 1911. In Seitz, Macrolepidoptera of the World 3 : 248, pi. 51. Stuttgart. 

1913. In Seitz, Macrolepidoptera of the World 11 : 313-315, pi. 28. Stuttgart. 



INDEX 



Synonyms in italics, page references to 
descriptions in bold type 

Adisura, 10, 70 

alarioides, 63, 64 

albicilia, 19, 70, 72 

albida, 11, 12, 93-94 

albipuncta, 18, 73, 74 

albirosea ssp., 16, 36, 37, 38-39 

albiseriata ssp., 16, 24, 25, 26, 27, 29 



arabica ssp., 20, 21 
artaxoides, 19, 81-82, 83, 84 

beatrix, 6, 11, 51, 54, 58-62, 62, 64, 66 
beatrix ssp., 14, 18, 58, 59-60, 61 
bechuana ssp., 17, 18, 39, 40, 45, 46-47 
belgaumensis , 79 
bimaculata, 85-91, 92 
bimaculata ssp., 18, 85, 86, 87, 90 
buchanani, 27 



INDEX 



99 



•alamaria, 84 

'anthylidia, 10 

;heesmanae, 11, 66, 69-70 

:heesmanae ssp., 14, 69-70 

hitinipyga, 6, 93 

hrvsita, 77, 78 

ontinnata, 30 

ornia ssp., 14, 19, 85, 86, 87, 88, 90 

ruentata, 11, 17, 39, 40, 47-49 

'uriihasa, 6 

langilc-nsis ssp., 14, 66, 69 

lecorata, 10, 24-29, 34, 36 

lecorata ssp., 16, 24, 25-26, 27, 29 

'epicta, 40, 43 

listicta, 11, 36-39 

listicta ssp., 11, 16, 36, 37, 38, 39 

lora, 18, 78-79 

pimethea, 4, 11, 12, 51, 54, 59, 61 62, 64, 66 



latinigra ssp., 14, 68-69 

latinigra — cheesmanae complex, 66-70 

Lecevfia, 6 

leucosticta, 10, 24, 29-33, 34, 36 

leucosticta ssp., 16, 29, 30-32, 33 

lineata, 65 



marginata, 47 
metaphaea, 19, 82, 83 
metarhoda ssp., 16, 24, 
mittoni, 18, 82-83 
modesta, 19, 82, 84 
multistriata, 55 



26, 27 29, 30 



nigrifasciata ssp., 17, 19, 85, 87, 90 
nigristriata, 59 
nigrolineata, 40, 43 
nigrolinc-ata form, 40 
nubila, 18, 76, 77-78, 79 
nuristana ssp., 10, 20, 2 1 



■««. 75. 7 6 
ssifascia, 10, 21-24 
iSBifascia ssp., 12, 21, 23-24 
'ana, 52 

aviceps, 11, 17, 39, 49, 50, 51 
avirosea ssp., 16, 36, 37, 39 
avistrigata, 11, 12, 14, 18, 51, 52, 54, 59, <>2, 
1 64, 65-66 
avocarnea, 10, 92 
unebris, 10, 16, 2(), 33, 36 
uscostriata ssp., 12, 17, 94, 95, 96 



pallescens, 87 

perstriata, 11, 94-96 

perstriata ssp., 18, 94, 95, 96 

philbyi, 10, 12, 19-21, 11 

pluritelifora ssp., 14, 19, 85, 87, 88, 90-91 

Pradatta, 6 

prochaskai, 4, 10, 16, 17, 34-35 

pulverulenta, 87 

quilengesi, 16, 18, 73, 74 



alatheae, 11, 39, 40-47, 49 

alatheae form, 40 

alatheae ssp., 17, 39. 40-45, 47 

alatheae — cruentata complex, 39-49 

lelicoverpa, 10 

ololeuca, 11, 16, 40, 50-51 

nitata, 40, 43 
nitata form, 40, 41 
rorata, 8 

liceyi ssp., 16, 29, 32, 33 



'.ncea, 4, 40, 45 
nceolata, 6, 40, 43 
tinigra, 1 1, 66-69 



radiata, 6, 8, 11, 51-54, 59, 62, 64, 66 
radiata ssp., 14, 51, 52, 54, 66 

rhodomelaleuca ssp., 12, 54, 55, 58 

rosacea, 11, 14, 51, 54, 59, 62, 64, 66 

rosastrigata, 90, 91 

rosea, 57, 58 

roseata ssp., 12, 22, 24 

roseivena, 11, 12, 51, 54, 59, 62, 63-64, 66 

rubristria, n, 51, 54-58, 59, 62, 64, 66 

rubristria ssp., 14, 54, 55, 58 



sanguistria, 55 
semifusca, 18, 79, 82 
senegalensis, 10 
splendens, 40, 43 
splendens form, 40 
striata, 93 
sublimis, 10, 19, 33, 35-36 



ioo INDEX 

tamburensis ssp., 14, 66, 70 uncta, 19, 70 

terracotta ssp., 14, 51, 52-54, 66 unifasciata ssp., 87, 88 

terracottoides, 12, 75-76 

Timora, 3, 4, 6, 7, 8, 10 vinula ssp., 4, 16, 29, 31, 32, 33 

tosta, 14, 18, 80 vittulata, 40, 43 

transvaalica ssp., 14, 54, 55, 57-58 

trifasciata ssp., 12, 58, 59, 60-61 zernytamsia ssp., 17, 94, 95, 96 



P. R. Seymour, M.Sc. 

Department of Entomology 

British Museum (Natural History) 

Cromwell Road 

London, SW7 5BD 



PLATE i 

Fig. 114. philbyi 6* (B.M. Neg. 471 18). 

Fig. 115. philbyi $ (B.M. Neg. 47120). 

Fig. 116. philbyi 6* (B.M. Neg. 471 19). 

Fig. 117. philbyi $ (B.M. Neg. 471 21). 

Fig. 118. fissifascia fissifascia 6" holotype (B.M. Neg. 43209). 

Fig. 119. fissifascia roseata $ paratype (B.M. Neg. 43208). 

Fig. 120. decorata decor ata $ lectotype (B.M. Neg. 43637)- 

Fig. 121. decorata decorata $ (B.M. Neg. 47130)- 

Fig. 122. decorata decorata ? (B.M. Neg. 47129). 

Fig. 123. decorata albiseriata 6" (B.M. Neg. 47133). 

Fig. 124. decorata metarhoda $ (B.M. Neg. 47131). 

Fig. 125. decorata metarhoda $ (B.M. Neg. 47132)- 



Hull. By. Mus. not. Hist. (Ent.) 27, 1 




I 



PLATE 1 



116 




U 

117 V 




120 




118 




121 




123 





124 



125 






PLATE 2 

Fig. 126. leucosticta leucosticta <J (B.M. Neg. 47125). 

Fig. 127. leucosticta leucosticta $ (B.M. Neg. 47123). 

Fig. 128. leucosticta leucosticta $ (B.M. Neg. 47122). 

Fig. 129. leucosticta vinula £ (B.M. Neg. 47126). 

Fig. 130. leucosticta joiceyi $ (B.M. Neg. 47127). 

Fig. 131. funebris <J holotype. 

Fig. 132. prochaskai $ (B.M. Neg. 46273). 

Fig. 133. sublimis <$ (B.M. Neg. 47180). 

Fig. 134. sublimis <$. 

Fig. 135. disticta albirosea $ lectotype. 

Fig. 136. disticta disticta £ holotype (B.M. Neg. 43630) 

Fig. 137. disticta flavirosea <J (B.M. Neg. 47096). 



Bull. Br. Mus. not. Hi si. (Ent.) 27, 1 



PLATE 2 



■ 








^^^^^^B 




&f 


Jffi**%^ 


p « 


'# 


1 


126 


? 127 


. * 


128 





131 




132 




133 




134 




w 




135 



136 



f 



137 



Fig. 138. 

Fig. 139. 

Fig. 140. 

Fig. 141. 

Fig. 142. 

Fig. 143. 

Fig. 144. 

Fig. 145. 

Fig. 146. 

Fig. 147. 

Fig. 148. 

Fig. 149. 



galatheae galatheae 
galatheae galatheae 



galatheae gaiatneae 
(B.M. Neg. 47106) 
galatheae galatheae 
\BM. Neg. 47105) 
galatheae galatheae 
galatheae galatheae 
(B.M. Neg. 47108) 
galatheae galatheae 
(B.M. Neg. 47107) 
galatheae galatheae 
galatheae bechuana 
galatheae bechuana 
galatheae bechuana 
galatheae bechuana 
galatheae bechuana 



PLATE 3 

form galatheae $ (Indian specimen) (B.M. Neg. 47104). 
form splendens $ (splendens lectotype) 

form galatheae $ (African specimen) 

form from Mackakos, Kenya <3 (B.M. Neg. 47109). 
form imitata $ (occurrence West & Central Africa) 

form nigrolineata $ (occurrence East & Central Africa) 

form from Luimbale, Angola $ (B.M. Neg. 471 10). 

9 paratype (B.M. Neg. 47112). 

$ paratype (B.M. Neg. 47119). 

$ holotype (B.M. Neg. 471 11). 

o* paratype (B.M. Neg. 471 14A). 

$ paratype (B.M. Neg. 471 14). 



Bull. Br. Mus. mil. Hut. (Ent.) 27, 1 



PLATE 3 








138 




141 




144 




139 




142 






145 




147 



148 




Fig. 150. 
Fig. 151. 
Fig. 152. 

Fig- 153- 
Fig. 154. 

Fig. 155- 
Fig. 156. 
Fig. 157. 
Fig. 158. 
Fig. 159. 
Fig. 160. 
Fig. 161. 



PLATE 4 

cruentata ? (B.M. Neg. 471 16). 

cruentata <$ (B.M. Neg. 47115). 

cruentata ? (B.M. Neg. 471 18). 

hololeuca $ (B.M. Neg. 47192). 

radiata radiata $ (B.M. Neg. 47138). 

radiata radiata £ lectotype (B.M. Neg. 43626). 

radiata terracotta $ (B.M. Neg. 43223). 

radiata terracotta $ lectotype (B.M. Neg. 43224). 

radiata terracotta $ (B.M. Neg. 51475). 

flaviceps £ (B.M. Neg. 47137). 

epimethea $ (B.M. Neg. 47193). 

epimethea <$ (B.M. Neg. 46274). 



Bull. Br. Mus. nat. Hist. (Ent.) 27, 1 



PLATE 4 








153 





f 






154 



155 



156 




158 




■:* 




1 



H 



% ^ 



159 



160 



161 



PLATE 5 

Fig. 162. ritbristria rubristria <J (specimen from Kete-Krachi, Ghana) (B.M. Neg. 47182) 
Fig. 163. rubristria rubristria £ (specimen from Fort Crampel, Central African Republic) 

(B.M. Neg. 47181). 
Fig. 164. rubristria rubristria $ (specimen from Kaolack, Senegal) (B.M. Neg. 47183). 
Fig. 165. rubristria rubristria $ (specimen from Navaro, Ghana) (B.M. Neg. 47184). 
Fig. 166. rubristria transvaalica $ lectotype (B.M. Neg. 43226). 
Fig. 167. rubristria transvaalica <$ (specimen from Njombe, Tanzania) 

(B.M. Neg. 47187). 
Fig. 168. rubristria transvaalica $ (specimen from Elisabethville, Congo (Kinshasa) 

(B.M. Neg. 47187). 
Fig. 169. rubristria transvaalica $ (specimen from Suna, Kenya) (B.M. Neg. 47186). 
Fig. 170. rubristria rhodomelaleuca $ (B.M. Neg. 47195 A). 
Fig. 171. terracottoides $ (B.M. Neg. 47147). 
Fig. 172. terracottoides $ (B.M. Neg. 47148). 
Fig. 173. terracottoides 6* holotype (B.M. Neg. 43210). 



iull. Br. Mus. nat. Hist. (Ent.) 27, 1 



PLATE 5 




168 



169 




./ 




171 



172 



173 



PLATE 6 



Fig. 174. Beatrix beatrix $ (B.M. Neg. 47184). 

Fig. 175. beatvix beatrix $ (B.M. Neg. 47190). 

Fig. 176. beatrix beatrix $ (B.M. Neg. 47191). 

Fig. 177. beatrix trifasciata $ (B.M. Neg. 47176). 

Fig. 178. beatrix trifasciata $ holotype (B.M. Neg. 47177). 

Fig. 179. rosacea £ lectotype (B.M. Neg. 47140). 

Fig. 180. rosacea $ (B.M. Neg. 47141). 

Fig. 181. roseivena $ holotype (B.M. Neg. 43228). 

Fig. 182. roseivena $ (B.M. Neg. 43229). 

Fig. 183. roseivena $ (B.M. Neg. 47134). 

Fig. 184. roseivena $ (B.M. Neg. 47135)- 

Fig. 185. semifusca $ holotype (B.M. Neg. 47174). 




PLATE 6 



176 




PLATE 7 



Fig. 


186 


Fig. 


187. 


Fig. 


188. 


Fig. 


189 


Fig. 


190. 


Fig. 


191. 


Fig. 


192. 


Fig. 


J93 


Fig. 


194 


Fig. 


195 


Fig. 


196. 


Fig. 


197. 



ftavistrigata q (lineata lectotype). 
flavistvigata £ (B.M. Neg. 47144). 
flavistrigata <$ (B.M. Neg. 47143). 
latinigra latinigra $ holotype (B.M. Neg. 43647). 
latinigra latinigra $ paratype (B.M. Neg. 47166). 
latinigra dangilensis $ holotype (B.M. Neg. 47167). 
cheesmanae cheesmanae J holotype (B.M. Neg. 47168). 
cheesmanae cheesmanae $ form paratype (B.M. Neg. 47195) 
cheesmanae tamburensis g holotype (B.M. Neg. 47169). 
uncta <$ lectotype (B.M. Neg. 47150). 
albicilia £ (B.M. Neg. 47179). 
albicilia $ (B.M. Neg. 47178). 



'//. Br. Mas. nat. Hist. (Hut.) 27, 1 




PLATE 7 





/ 


_^„ 
















188 





f*ft 






192 



193 



194 







? 





195 



196 



197 



Fig. 


198 


Fig. 


199 


Fig. 


200 


Fig. 


201 


Fig. 


202 


Fig. 


203 


Fig. 


204 


Fig. 


205 


Fig. 


206 


Fig. 


207 


Fig. 


208 


Fig. 


209 



PLATE 8 

albipuncta $ lectotype (B.M. Neg. 43644). 

albipuncta ? (B.M. Neg. 47188). 

quilengesi $ holotype (B.M. Neg. 47101). 

quilengesi ? paratype (B.M. Neg. 47102). 

quilengesi <$ paratvpe (form from Okahanja, South West Africa) 

(B.M. Neg. 47103)- 

nitbila $ (B.M. Neg. 47170). 

nubila $ (B.M. Neg. 47171). 

nubila $ (chrysita lectotype). 

dor a $ lectotype (B.M. Neg. 47145). 

dora $ (B.M. Neg. 47146). 

tosta ? (B.M. Neg. 47172). 

tosta $ lectotype (B.M. Neg. 47173). 



till. Br. Mus. nat. His/. (Ent.) 27, 1 



PLATE 8 



■ 



* 



198 



199 





W 



201 



202 



! 




205 





3 



VI 



207 



208 



200 




203 




206 




209 






PLATE 9 



Fig. 210. 
Fig. 211. 
Fig. 212. 
Fig. 213. 
Fig. 214. 
Fig. 215. 
Fig. 216. 
Fig. 217. 
Fig. 218. 
Fig. 219. 
Fig. 220. 
Fig. 221. 
Fig. 222. 



bimacitlata 
bimaculata 
bimaculata 
bimaculata 
bimaculata 
bimaculata 
bimaculata 
bimaculata 
bimaculata 
bimaculata 
bimaculata 
bimaculata 
flavocamea 



bimaculata $ holotype (B.M. Neg. 47151 
bimaculata $ (B.M. Neg. 47152). 
cornia $ paratype (B.M. Neg. 47155). 
cornia 9 paratype (B.M. Neg. 47154). 
cornia <$ paratype (B.M. Neg. 47153). 
cornia $ holotype (B.M. Neg. 47156). 
nigrifasciata £ (intergrade) (B.M. Neg. 



nigrifasciata $ 
nigrifasciata $ 
nigrifasciata $ 
pluritelifora $ 
pluritelifora $ 



intergrade) (B.M. 
(B.M. Neg. 47160), 
(B.M. Neg. 47159). 
(B.M. Neg. 47158). 
(B.M. Neg. 47157)- 



Neg. 



48161) 
47162) 



$ holotype (B.M. Neg. 47149). 



nil. Br. Mus. nat. Hist. (Ent.) 27, 1 



PLATE 9 



210 





PLATE 10 



Fig. 223. albida $ {chitinipyga lectotype). 

Fig. 224. albida $ (B.M. Neg. 47098). 

Fig. 225. albida $ (B.M. Neg. 47100). 

Fig. 226. per striata fuscostriata $ paralectotype. 

Fig. 227. perstriata fuscostriata $ paralectotype. 

Fig. 228. perstriata fuscostriata $ (from type-series) 

Fig. 229. perstriata perstriata <$ holotype (B.M. Neg. 47163). 

Fig. 230. perstriata zemytamsia $ (B.M. Neg. 47165). 

Fig. 231. artaxoides $ (B.M. Neg. 47175). 

Fig. 232. metaphaea ? (B.M. Neg. 47136). 

Fig. 233. mittoni $ lectotype (B.M. Neg. 43215). 

Fig. 234. modesta $ {calamaria lectotype) (B.M. Xeg. 43652) 



nil. Br. Mits. nat. Hist. (Ent.) 27, 1 



PLATE 10 



225 




228 



229 




) 



232 



230 



233 



231 




234 



^nSHSN 



A LIST OF SUPPLEMENTS 

TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 
18 plates, 270 text-figures. August, 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 : 500 text-figures. September, 

1965- £3.25. 

5. Ahmad, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156 : 
475 text-figures. November, 1965. (out of print) £2.15. 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. 

8. Fletcher, D. S. A revision of the Ethiopian species and a check fist of the 
world species of Chora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 
text-figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera). Pp. 509. August, 1967. £8.50. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322: 348 text-figures. August, 1967. £8. 

11. Mound, L. A. A review of R. S. .Bagnall's Thysanoptera Collections. Pp. 172: 

82 text-figures. May, 1968. £4 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. 
November, 1968. £5. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
.Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North-Western Europe 
(Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 
68 plates, 15 text-figures. October, 1971. £12. 



•jjft; Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 



A, TAXONOMIC REVIEW OF THE 
?/ SPECIES OF CINARA CURTIS 
F^ OCCURRING IN BRITAIN 
■ (HEMIPTERA : APHIDIDAE) 



V. F. EASTOP 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 27 No. 2 

LONDON : 1972 



A TAXONOMIC REVIEW OF THE SPECIES OF 

CINARA CURTIS OCCURRING IN BRITAIN 

(HEMIPTERA: APHIDIDAE) 




BY 

VICTOR FRANK EASTOP 






Pp. 101-186 ; 41 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 27 No. 2 

LONDON: 1972 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history) instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 27 No. 2 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. not. Hist. (Ent. 



Trustees of the British Museum (Natural History), 1972 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



Issued 14 March, 1972 Price £3.40 



A TAXONOMIC REVIEW OF THE SPECIES OF 

CINAR CURTIS OCCURRING IN BRITAIN 

(HEMIPTERA: APHIDIDAE) 

By V. F. EASTOP 

CONTENTS 

Page 

Synopsis ........... 103 

Introduction .......... 103 

Nomenclature .......... 104 

Generic Synonymy. ......... 104 

Systematics ........... 106 

Generic Diagnosis ......... 107 

Taxonomy ........... 108 

Biology ........... 109 

Key to the apterae viviparae of the British species of Cinara . no 

Key to the alatae viviparae of the British species of Cinara . 115 

The British species of Cinara . ....... 123 

Host-plants of the British species of Cinara . . . . 170 

Acknowledgements ......... 171 

blometric data for the apterae viviparae of the british species of 

Cinara ......... following p. 172 

BlOMETRIC DATA FOR THE ALATAE VIVIPARAE OF THE BRITISH SPECIES OF 

Cinara ......... following p. 172 

Discriminants for the apterae viviparae of the British species of 

Cinara ........... 173 

Discriminants for the alatae viviparae of the British species of 

Cinara ........... 174 

BlOMETRIC AND OTHER DATA FOR THE SEXUALES OF THE BRITISH SPECIES 

of Cinara . . . - . . . . . . . 175 

References . . . . . . . . . . .176 

Index ............ 184 

SYNOPSIS 

Keys are given for the identification of the viviparous morphs of the 21 species of Cinara 
known from Britain. Biometric data for each species are tabulated and a summary is given of 
their known host-plant range, geographical distribution and biology. Six specific synonyms 
are newly established and one former variety (stroyani Pasek) is raised in status to species. 



INTRODUCTION 

The purpose of this paper is to provide a means of identifying the species of Cinara 
occurring in Britain. As understood here, Cinara includes Cupressobium and 
contains about 200 described species living on Coniferae of the families Pinaceae 
and Cupressaceae. About 150 of the species are described from North America, 
20 from Japan and the Oriental region and 30 are European or Mediterranean in 
origin. Twenty-one species have been seen from Britain, but not more than five 
of these can be native since the other 16 species live on introduced host-plants. 



io 4 V. F. EASTOP 

Most of the introduced aphids are from central Europe and British material fits 
central European descriptions. There appears to be another fauna south of the 
Alps, as it is difficult to relate British material to the descriptions of the Italian 
fauna by Del Guercio (1909) and to the Spanish fauna by Gomez Menor (1962). 

Recent accounts of the species of Cinara from various parts of Europe (Pasek, 
1952-1954; Heinze, 1962; Szelegiewicz, 1962; Pintera, 1966) have made it much 
easier to identify the British species. Recent authors have made great use of the 
chaetotaxy of the processus terminalis of the last antennal segment and the shape 
of the first segment of the hind tarsus. As the last antennal segments and hind 
legs of trapped alatae of Cinara are often lost and as no previous account includes 
all the British species, this paper started as an attempt to construct a key to alatae 
using characters on less deciduous structures, such as the rostrum and abdomen. 

The principal recent works on the taxonomy of Cinara include Braun, 1938 : 
461-510, Borner, 1949 : 52-60, Heinze, 1962 : 145-178, Central Europe; Pasek, 1954 : 
1-319 and Pintera, 1966 : 281-321, Czechoslovakia; Szelegiewicz, 1962 : 63-98, 
Poland; Shaposhnikov, 1964:521-524, European U.S.S.R. ; Bodenheimer & 
Swirski, 1953 : 182-183, 245-246, Middle East; Narzikulov, 1962 : 111-118, Central 
Asia; Inouye, 1970 : 57-102, Japan; Paik, 1965 : 16-22, Korea; Takahashi, 1931 : 
22-24, Taiwan; Blanchard, 1939 : 859-870, Argentina; Tissot, 1939 : 34-47, Florida; 
Palmer, 1952 : 20-52, Rocky Mountains. Hottes has published many papers on 
North American Cinara between 1930 and 1965. G. A. Bradley, 1956-1968, has 
published a series of papers on Canadian Cinara but his monograph has not yet 
appeared. Eastop, 1961 : 74-75 & 1966 : 525-529 gave keys to the species of 
Cinara introduced to Africa and Australia respectively. 

NOMENCLATURE 

There has been confusion about the use of Lachnus Burmeister and Cinara Curtis 
because both were described in 1835 and there was doubt about the identity of the 
type-species of both genera. As a result various replacement names were proposed 
but subsequently proved unnecessary. Hottes (1930 : 185-188) and Theobald 
(1929 : 352) discuss the usage of Cinara and Lachnus. 

Although Aphis pini L. has been accepted as the type-species of Cinara for the 
last 40 years, there has been a dispute as to the species of Cinara to which the 
name pini L. should be applied. Stroyan (1955 : 332-333) summarizes the position. 

GENERIC SYNONYMY 

CINARA Curtis, 1835 

[Aphis L., partim, auctores diversi, 1758-1852.] 
[Lachnus Burmeister, partim, auctores diversi, 1 835-1933.] 
*Cinara Curtis, 1835 : part 576. Type species: Aphis pini L., 1758 : 453. 
*Lachniella Del Guercio, 1909 : 286. Type-species: Lachnus fasciatus Burmeister, 1835 = 

costata Zetterstedt, 1828. 
*Todolachnus Matsumura, 1917 : 381. Type-species: Todolachnus abietis Matsumura, 1917 : 
381-382. 
Wilsonia Baker, 1919a : 212, nee Bonaparte, 1838. Type-species: Lachniella gracilis Wilson, 
1919 : 20-21. 



REVIEW OF CINARA IN BRITAIN 105 

Dilachnus Baker, 19196 : 253, nee Fairmaire, 1896. Type-species: Lachniella gracilis Wilson, 

1919 : 20-21. 
Panimerus Laing in Theobald, 1929 : 129, nee Eaton, 1913. Type-species: Lachniella gracilis 

Wilson, 1919 : 20-21. 
*Neochmosis Laing in Theobald, 1929 : 129, footnote. Type-species: Lachniella gracilis 

Wilson, 1929 : 20-21. 
Neodimosis Toth, 1935 : 495. [Incorrect subsequent spelling of Neochmosis Laing.] 
*Cinaria Bonier, 1939 : 76. Type-species: Aphis laricis Walker. 
*Cinarina Borner, 1939 : 76. Type-species: Lachnus viridescens Cholodkovsky = bogdanowi 

Mordwilko. 
*Cinaropsis Borner, 1939 : 76. Type-species: Lachnus pinicola Kaltenbach, 1843, sensu 

Borner = pilicornis Hartig. 
*Dinolachnus Borner, 1940 : 1. Type-species: Lachniella cilicica var. cecconii Del Guercio = 

abieticola Cholodkovsky. 
*Cupressobium Borner, 1940 : 1. Type-species: Aphis juniperi De Geer. 
*Cinarella Hille Ris Lambers, 1948 : 275 [as subgenus of Cinara Curtis]. Type-species: 

Lachnus pineus Mordwilko. 
*Subcinara Borner, 1949 : 59 [as subgenus of Cinara Curtis]. Type-species: Cinara braani 

Borner. 
Cinavella Borner, 1949 : 59, nee Hille Ris Lambers, 1948 [as subgenus of Cinara Curtis]. 

Type-species: Cinara laricicola Borner, 1939 nee Matsumura, 1917 = boerneri Hille Ris 

Lambers. 
*Laricaria Borner, 1949 : 59 [as subgenus of Cinaria Borner]. Type-species: Cinara kochiana 

Borner. 
Pityaria Borner, 1949 : 59 [as subgenus of Cinaria Borner]. Type-species: ' Lachnus pruinosus 

Htg. = bogdanowi Md.' 
*Mecinaria Borner, 1949 : 59 [as subgenus of Cinaria Borner]. Type-species : Aphis piceae 

Panzer. 
*Cinarellia Borner, 1951 : ?; 1952 : 41. Type-species: Cinara laricicola Borner, 1939 (nee 

Matsumura, 1917) = boerneri Hille Ris Lambers. Borner, 1952 : 41 lists Cinarellia Borner, 

1951 with Cinarella Borner, 1949 nee Hille Ris Lambers, 1948, as a synonym but gives no 

reference to the 1951 paper and I have not been able to find it. 
*Buchneria Borner, 1952 : 41, 242. Type-species: Aphis pectinalae Nordlinger. 
[Eulachnus Del Guercio, Borner, 1952 : 241. Borner considered Eulachnus mingazzinii Del 

Guercio (= piniphila Ratzeburg) to be the type-species of Eulachnus Del Guercio. How- 
ever, the International Commission on Zoological Nomenclature has since ruled that the 

type-species is Lachnus agilis Kaltenbach (Bull. zool. Nom. 22 : 188-189, J 965), and so 

Eulachnus is not a synonym of Cinara and is available in its most widely used sense with 

Protolachnus Theobald as a synonym] 
Neocinaria Pasek ms [as subgenus of Cinaria Borner]. Type-species: Cinara escherichi 

Borner [see Pinteia, 1966 : 281-282]. 
Pseudocinara Pasek, ms [as subgenus of Cinara Curtis]. Type-species: Lachnus neubergi 

Arnhardt [see Pintera, 1966 : 282]. 

The names marked with an asterisk (*) are available for subgenera. 

Alphabetical list of the type-species of the subgenera and synonyms of Cinara. 

abieticola Cholodkovsky, 1899. Type of Dinolachnus Borner, 1940 (as cilicica var. 

cecconii) . 
abietis Matsumura, 1917. Type of Todolachnus Matsumura, 1917. 
boerneri Hille Ris Lambers, 1956. Type of Cinarella Borner, 1949 (nee Cinarella 

Hille Ris Lambers, 1948) and Cinarellia Borner, 1951/1952, as laricicola Borner, 

1939 nee Matsumura, 1917. 



io6 V. F. EASTOP 

bogdanowi Mordwilko, 1895. Type of Cinarina Borner, 1939 (as viridescens) and 

Pity aria Borner, 1949 (as pruinosus) . 
brauni Borner, 1940. Type of Subcinara Borner, 1949. 
cilicica var. cecconii Del Guercio = abieticola Cholodkovsky. 
costata Zetterstedt, 1828. Type of Lachniella Del Guercio, 1909 (as fasciata) . 
escherichi Borner, 1950. Type of Neocinara Pasek, m.s. 
fasciata Burmeister, sensu Del Guercio, 1909 = costata Zetterstedt. 
gracilis Wilson, 1919. Type of Wilsonia Baker, 1919 (nee Wilsonia Bonaparte, 

1838 etc.) ; Dilachnus Baker, 1919 (nee Fairmaire, 1896) ; Panimerus Laing, 1929 

(nee Eaton, 1913) and Neochmosis Laing, 1929. 
juniperi De Geer, 1773. Type of Cupressobium Borner, 1940. 
kochiana Borner, 1939. Type of Laricaria Borner, 1949. 
laricicola Borner, 1939 nee Matsumura, 1917 = boerneri Hille Ris Lambers. 
laricis Walker, 1848. Type of Cinaria Borner, 1939. 
neubergi Arnhardt, 1930. Type of Pseudocinara Pasek, m.s. 
pectinatae Nordlinger, 1880. Type of Buchneria Borner, 1952. 
piceae Panzer, 1801. Type of Mecinaria Borner, 1949. 
pilicornis Hartig, 1841. Type of Cinaropsis Borner, 1939 (as pinicola) . 
pineus Mordwilko, 1895. Type of Cinarella Hille Ris Lambers, 1948. 
pini L., 1758. Type of Cinara Curtis, 1835. 
pinicola Kaltenbach, 1843 sensu Borner = pilicornis Hartig. 
pruinosus Hartig, 1841 sensu Borner = bogdanowi Mordwilko. 
viridescens Cholodkovsky, 1898 = bogdanowi Mordwilko. 

SYSTEMATICS 

The European species of Cinara fall into fairly well defined groups which have 
been dignified as genera and subgenera. The British species could be arranged as 
in the table on page 175. Although there are evident groups of species when the 
world fauna is considered, I have not been able to find characters to separate them 
absolutely. For instance, Cinara konoi Inouye, C. longipennis (Matsumura) and 
C. todocolus Inouye from the Far East, C. chinookiana Hottes, C. lasiocarpae 
(Gillette & Palmer) and C. sonata Hottes from America all resemble C. abieticola 
(Cholodkovsky) and live on Abies, suggesting that Dinolachnus should be regarded 
as at least subgenerically distinct. Cinara sonata, however, has four sub-apical 
setae on the processus terminalis and a rim around the primary rhinaria, just the 
characters distinguishing Cinara s. str. from the other Dinolachnus. Cinara 
{Lachniella) costata (Zetterstedt) from Picea in the palaearctic region has character- 
istically pigmented wings and short first tarsal segments, as does C. (L.) comata 
Doncaster from the Himalayas. The dorsal length of the first tarsal segments 
of their hind legs is only about 0-6-0-8 of the basal diameter, and about 0-25-0-33 of 
the ventral length of the segment. In C. (L.) nimbata Hottes from Picea engelmanii 
in North America the dorsal length of the first tarsal segments of the hind leg is 
about 1-2-1-5 times as long as its basal diameter and about 0-4 of its ventral length. 
Apart from bearing rather shorter hairs, C. (L.) nimbata is otherwise similar to 
C. (L.) costata. Cinara difficilis Hottes from Juniperus in North America has short 



REVIEW OF CINARA IN BRITAIN 107 

first tarsal segments and long fine hairs like a Cupressobium, but has a a rim round 
the primary rhinaria and bears five sub-apical setae on the processus terminalis. 
C. manitobensis Bradley from Junipems in North America has four sub-apical setae 
on the processus terminalis, a rim round the primary rhinarium and the appearance 
of a Cinaropsis. Cinara coloradensis (Gillette) and C. hottest Gillette & Palmer 
live on Picea in North America but resemble the C. pint group (= Cinara s. str.) 
which are confined to Larix and Pinus in Europe. I have not been able to find any 
satisfactory subgeneric separation for the world fauna. 

Heie (1967 & 1969) gives a comprehensive account of the aphids known from 
baltic amber, which is thought to be the petrified resin of Pinites succinifera Goeppert, 
an extinct conifer related to Pinus. Heie studied 103 aphids from amber without 
finding a single member of the subfamily Lachninae to which Cinara belongs. 
Pinites resin may have been more sticky in the rather warmer conditions of the 
lower Oligocene than Pinus resin is today. The absence of Cinara from present-day 
resin on English conifers may be due to resin being most sticky in high summer 
when Cinara are little evident, being either subterranean or perhaps in reproductive 
diapause. I have not been able to find insects trapped in resin on living conifers 
in order to compare them with the aphid fauna of the tree. Under English con- 
ditions, conifer resin usually occurs only in small quantities, and the surface is not 
sticky. The large size, complete wing venation, dense pubescence, trace of a third 
tarsal segment, distinct 4th and 5th rostral segments, compound eyes in all forms, 
absence of host-plant alternation and association with coniferae are characters 
suggesting that Cinara belongs to an old group of aphids. The absence of Lachninae 
from baltic amber, however, and the short straight radius arising from near the tip 
of the stigma of the fore wing suggest that the group is more recent. 

GENERIC DIAGNOSIS 

Medium-sized to large aphids, body 2-8 mm long. General appearance as in 
Text-fig. 28 (p. 154). Yellowish, red-brown, dark brown or sometimes green 
aphids. Antennal tubercles absent. Antennae 6-segmented, 0-2-0-6 as long as 
the body. Processus terminalis short, 0-08-0-33 as l° n g as * ne base of the sixth 
antennal segment and bearing 3 apical and 2-1 1 sub-apical setae. Apterae 
usually with a rhinarium at the apex of the fourth antennal segment and often also 
with a secondary rhinarium on the fifth segment just basadof the primary rhinarium. 
Alatae viviparae with secondary rhinaria distributed: III, 1-18; IV, 0-6; V, 0-4; 
VI, 0. Antennae, body and legs often covered with numerous long fine hairs 
(Text-figs 1, 19, 28-31, 38-39) or the hairs may be shorter and only about equal in 
length to the diameter of the third antennal segment (Text-figs 6, 18, 22). Rostrum 
evidently consisting of five segments, the suture between segments 4 and 5 distinct 
(Text-figs 4, 9, 10, 25, 36). The fourth rostral segment bears from 2 to 60 accessory 
hairs but most species bear 4-14 accessory hairs arranged in two rows bordering 
the groove for the stylets. Wing venation as in Text-figs 13, 20, 37; the radius 
originates from the end of the pterostigma and extends in a straight line to the wing 
apex. Hind wing with two oblique veins. Tarsi 2-segmented but when the basal 
segment is long as in the subgenus Cinarella, it often bears traces of a suture suggest- 



io8 V. F. EASTOP 

ing that it is composed of two fused segments. The first tarsal segment bears nine 
or more ventral hairs and the second segment bears eight or more long dorsal hairs 
and about 16 shorter ventral hairs. Empodial hairs very short, only about o-i times 
as long as the claws. Mid thoracic furca of apterae with a short base. 

Siphunculi placed on pigmented cones whose diameter varies with the degree 
of pigmentation but is often about equal to the length of the third antennal segment. 
Cauda crescent-shaped and bearing about 20 hairs. Eighth abdominal tergite 
bearing y-yy hairs. 

TAXONOMY 

The ratios of the fifth antennal segment to the other antennal segments have 
been used to recognize species. The fifth antennal segment tends to increase in 
size more with increased body size than does the sixth antennal segment. Thus 
the antennal V : VI ratio tends to increase with body size and the differences be- 
tween large and small specimens are greatest in species with the greatest differences 
between the lengths of the segments. In Cinara laricis and C. piceae with a rela- 
tively long antennal V, the V : VI ratio of large specimens varies from 1-9-2-4 and 
for C. abieticola from 1-9-2-2, while the ratio varies from 1-2-1-5 in small specimens 
of all three species. In species where antennal VI is as long as or longer than V 
(Cupressobium, Lachniella, Buchneria), V and VI increase in length more or less 
in unison and in proportion to the body length. These species have an antennal 
V : VI ratio of 0-7-1-1 over their whole size range. In species with antennal V 
only a little longer than VI, such as C. pini, C. bogdanowi, C. pilicornis (i.e. i Cinara 
s. str. and Cinaropsis) , the large specimens have the antennal V : VI ratio varying 
from 1-3-1-8 and the small specimens from 1-0-1-4. 

The adults, particularly of fundatrices and oviparae, may look similar to fourth 
instar larvae. The adult form can be recognized by the shape of the sub-genital 
plate. The rudimentary gonopophyses are also a sign of maturity but are sometimes 
difficult to detect in adults and may sometimes be detected before the final moult. 

Fundatrices usually have shorter antennae and legs than the later generations and 
the processus terminalis is particularly short. The fundatrices often bear more 
abdominal hairs than later generations but sometimes bear fewer hairs on the second 
and sixth antennal segments than later generations. Cinara piceae is unusual 
in that the fundatrices tend to bear fewer abdominal hairs as well as fewer hairs 
on the fourth rostral segment and second antennal segment than in the summer 
generations. Cinara piceae is also unusual in that while oviparae are common, 
males are unknown. The fundatrices of C. piceae may arise from unfertilized 
eggs or be fundatrices spuriae derived from an as yet undetected overwintering 
larva. The apterous summer generations of some species bear a well developed 
mesosternal tubercle (Text-fig. 11, p. 127) but the tubercle is absent or only weakly 
developed in the spring generations of these species. 

The eighth abdominal tergite of the oviparae may be pale but in Cinara boerneri it 
bears a small amount of pigmentation similar to the apterae viviparae. In the 
viviparae of most species of Cinara the eighth abdominal tergite is well pigmented. 
The sub-genital plate of the oviparae is large and densely hairy and there is often 



REVIEW OF CINARA IN BRITAIN 109 

a group of latero-ventral hairs on either side of the eighth abdominal tergite. The 
hind tibiae of the oviparae are slightly thickened and bear numerous pseudosensoria 
in the species with alate males. The hind tibiae of the oviparae of C. acutirostris, 
which has apterous males, are devoid of pseudosensoria Samples of oviparae of 
C. boerneri collected together with males show evident pseudosensoria but in one 
sample in which only oviparae were collected the pseudosensoria are indistinct. In 
oviparae of C. kochiana the pseudosensoria are present but indistinct even when 
males are present. 

The male genitalia of Cinara boerneri (Text-figs 14-16), C. cupressi (Text-fig. 21) 
and of C. pectinatae (Text-fig. 26) may be characteristic for each species. The 
problem when using male genitalia for aphid taxonomy is that, being soft, prepara- 
tions from the same sample (Text-figs 14-16) may look different merely through 
lying in different positions. 

Descriptions of each species are not given individually but biometric data for 
apterae and alatae viviparae and discriminants for apterae and alatae viviparae 
are given in the tables between pages 172-173. Data for sexuales are given in the 
table on page 175. 

BIOLOGY 

As far as is known Cinara species live only on Coniferae and usually only on 
Pinaceae and Cupressaceae. Most of the species living on Pinaceae are specific 
to one or to a few closely related species of Abies, Larix, Picea or Finns. No species 
are known which live on more than one of these genera although some species nor- 
mally living on Abies can breed successfully on Cedrus. Some Cinara living on 
Cupressaceae have a wider host-plant range, occurring on species in several genera 
of conifers although often not on all the species in these genera. For instance the 
usual hosts of Cinara cupressi are Cupressus macrocarpus, Thuja occidentalis and 
Juniperus virginiana. This host-plant range may only be a reflection of the lower 
level of agreement among botanists of the generic classification of Cupressaceae 
compared with the Pinaceae. Cinara species feed on the bark of their host-plants 
and some species are associated with lesions of Pine rust fungi (Tissot & Pepper, 
1967 : 1-10). This association with rust fungi may be analogous with the habit 
of feeding on other insects galls found in other aphids in various systematic groups. 
The presence of the fungi may act as a physiological sink and like galls, stimulate 
the translocation of the metabolic products on which the aphids feed. 

Most species of Cinara overwinter in the egg-stage from which an apterous 
fundatrix develops in the spring. The second or third generation is commonly 
winged and the summer may be spent on the roots of the same species of conifer 
on which the overwintering egg was laid. The remaining generations may be 
wingless or some alatae may occur from July onwards. In the autumn oviparae 
and winged or sometimes wingless males are produced. Cinara piceae is a widely 
distributed species in which oviparae have often been found but males are unknown. 
Some species of Cinara continue to reproduce on the aerial parts of their host all 
through the summer. Others remain on the aerial parts but adult apterae collected 
in late July or August may not contain fully developed embryos, which suggests 



no V. F. EASTOP 

that reproductive diapause occurs. Many species of Cinara are associated with 
ants (Way, 1963 : 307-344). Several species of Cinara are of importance to bee- 
keepers in Central Europe as their honeydew is the source of 'forest honey'. 

Key to the apterae viviparae of the British species of Cinara 

1 Fourth rostral segment (Text-fig. 25) 330-420(1 long, 2 -4-3 o times as long as the 

fifth rostral segment and bearing 24-32 accessory hairs arranged in 4 longi- 
tudinal rows of 6-8 hairs per row. Second antennal segment bearing 21-31 
hairs. Base of sixth antennal segment (Text-fig. 23) bearing 15-24 hairs. 
Processus terminalis (Text-fig. 24) bearing 6-9 sub-apical setae. Mesosternal 
tubercle moderately developed. Second segment of hind tarsus 1-0-1-2 
times as long as the fourth rostral segment. Fifth abdominal tergite bearing 
18-30 hairs between the siphunculi. Longest hairs on the third abdominal 
tergite 15-25(1 long. On Larix ..... kochiana (p. 143) 

- Fourth rostral segment 120-370(1 long, 1 -5-3-0 times as long as the fifth rostral 

segment and bearing 2-17 accessory hairs arranged in two longitudinal rows 
of 1-9 hairs per row. (Text-figs 4, 9, 10). Second antennal segment bearing 
5-26 hairs. Base of the sixth antennal segment bearing 4-20 hairs and 
processus terminalis usually with only 3 or 4 sub-apical setae but sometimes 
with 5— 11. If either the second antennal segment bears 20 or more hairs 01 
the base of the sixth antennal segment bears 19 or more hairs (abieticola, 
bogdanowi, stroyani), then the fifth abdominal tergite bears 40-95 hairs 
between the siphunculi and the longest hairs on the third abdominal tergite 
are 60-27051 long. If the processus terminalia bears 5-1 1 subapical setae, 
then either the mesosternal tubercle is absent and the second antennal 
segment, base of the sixth antennal segment, fourth rostral segment, fifth 
abdominal segment between the siphunculi and the eighth abdominal 
tergite each bear only 5-17 hairs (piceae, escherichi) or if the fifth abdominal 
tergite bears 23-36 hairs between the siphunculi (brauni), then abdomi- 
nal tergites 5-7 are fused in a solid dark patch ..... 2 

2 (1) Abdominal tergites 5-7 bearing a solid dark patch encompassing the siphun- 

cular cones. 

Processus terminalis 50-100(1 long, 22-33% OI the total length of the sixth 
antennal segment and bearing 5-7 sub-apical setae. Second segment of the 
hind tarsus 360-4430(1 long, 1-1-1-5 times as long as the fourth rostral seg- 
ment which is 250-320(1 long. Eighth abdominal tergite bearing only 
10-15 hairs, the longest of which are 140-180(1 long. Longest hair on the 
third abdominal tergite 140-190(1 long. Third antennal segment (Text-fig. 
19) 500-700(1 long and 4-5-9-5 times as long as the longest hair, 70-120(1, 
borne on it. On Pinus nigra . ...... brauni (p. J33) 

- Pigmentation absent from tergites 5 and 6 or present only as isolated scleroites, 

any pigmentation present on each tergite being distinct from that on the 
other tergites. 

Processus terminalis usually bearing only 3 or 4 sub-apical setae but if 
with 5-1 1 then the longest hair on the third abdominal tergite is 5-30(1 long 3 

3 (2) Some at least of the hairs on the anterior abdominal tergites arising from 

scleroites with a diameter several times that of the hair base. 

Medium to large aphids, body 2-5-5-2 mm long, hind tibiae 1-5-3-5 mm 
long. Second antennal segment bearing only 5-10 hairs, base of the sixth 
antennal segment bearing 2-8 hairs. Fourth rostral segment bearing 4-8 
accessory hairs. First segment of hind tarsus 2-9-5-7 times as long as its 
own basal width. Third, fourth and fifth antennal segments often without 
secondary rhinaria. On A bies, Larix and Pinus ..... 4 






REVIEW OF CINARA IN BRITAIN in 

Scleroites absent or if present their diameter is less than 2-5 times that of the 

hair base ............ 7 

(3) Mesosternal tubercle (Text-fig. 11) present in the summer generations but 

sometimes only weakly developed or absent in the spring generations. 
Sixth antennal segment 100-160+ 22-3710 long. Ultimate rostral segment 
150-210+ 8o-ioo[i long. Second segment of hind tarsus 270-390(0. long, 
2-0-2-5 times as long as the first segment of the hind tarsus which is iio-i8o[o 
long and is 2 -9-4-1 times as long as its own basal diameter. Fourth antennal 
segment often and fifth antennal segment usually bearing a secondary 
rhinarium. On Larix and Pinns ........ 5 

Mesosternal tubercle absent. Sixth antennal segment 140-230+ 42-71(0 long. 
Ultimate rostral segment 210-290+ 1 10-170(0, long. Second segment of 
hind tarsus 350-530(0. long, 1-4-2-0 times as long as the first segment of the 
hind tarsus which is 190-330(0 long and 3 -7-5-7 times as long as its own basal 
diameter. Third and fourth antennal segments usually and fifth often 
without secondary rhinaria. On Abies and Pinus ..... 6 

(4) Third antennal segment 6-5-13 times as long as the longest hair, 40-80(0, 

borne on it. Hind tibiae 1-5-2-9 mm long, 17-34 times as long as the longest 
hairs, 65-125(0 borne on them. Fourth rostral segment 160-210U. long. 
Third antennal segment usually without a rhinarium, fourth antennal seg- 
ment often without a rhinarium. Base of the sixth antennal segment bearing 
4-7 hairs. Fifth abdominal tergite bearing 20-42 hairs between the 
siphunculi. Eighth abdominal tergite bearing 13-23 hairs. On Larix. 

laricis (p. 146) 
Third antennal segment 4-3-6-0 times as long as the longest hair, 110-150(0, 
borne on it. Hind tibiae 2 -5-2 -7 mm long, 14-16 times as long as the longest 
hairs, 170-190(0. borne on them. Fourth rostral segment 150-160(0 long. 
Third and fourth antennal segments each usually bearing a rhinarium. 
Base of the sixth antennal segment bearing 8-10 hairs. Fifth abdominal 
tergite bearing 40-50 hairs between the siphunculi. Eighth abdominal 
tergite bearing 20-24 hairs. On Pinus sylvestris . . pinihabitans (p. 162) 

(4) Siphuncular cones (Text-fig. 33) 250-700(0 in diameter. Third antennal seg- 
ment 490-740(0 long and 3-0-6-5 times as long as the longest hair, 90-210(0, 
borne on it. Fourth rostral segment 210-290(0. long, 1 -6-2-2 times as long as 
the fifth rostral segment, which is 110-170(0 long. Hind tibia 1-8-3-4 mm 
long and 12-20 times as long as the longest hair, 120-230(0, borne on it. 
Longest hair on the third abdominal tergite 95-210(0, on the eighth abdominal 
tergite 130-200(0 long. Eighth abdominal tergite bearing 23-28 hairs. 
Green aphid with paired dorsal longitudinal yellow stripes. On Abies. 

pectinatae (p. 148) 
Siphuncular cones (Text -fig. 33) 250-700(0 in diameter. Third antennal segment 
490-740(0 long and 3-0-6-5 times as long as the longest hair, 90-210(0, borne 
on it. Fourth rostral segment 210-290(0 long, 1-6-2-2 times as long as the 
fifth rostral segment, which is 110-170(0. long. Hind tibia 1-8-3-4 mm l° n g 
and 12-20 times as long as the longest hair, 120-230(0, borne on it. Longest 
hair on the third abdominal tergite 95-210(0, on the eighth abdominal 
tergite 120-230(0. long. Eighth abdominal tergite bearing 13-26 hairs. 
Yellowish brown aphids bearing numerous darker brown spots. On Pinus. 

pinea (p. 156) 

(3) Eighth abdominal tergite bearing 7-18 hairs. Fourth rostral segment 300-370(0 
long. Fifth abdominal tergite bearing 6-17 hairs between the siphunculi. 
Longest hairs on the third abdominal tergite 5-26(0 and on the eighth 
abdominal tergite 45-130(0. Processus terminalis bearing 6-1 1 sub-apical 
setae. 



ii2 V. F. EASTOP 

Body length 3-2-67 mm. Hind tibiae 2-4-5-2 mm long and 36-84 times 
as long as the longest hair, 50-900, borne on them. Sub-genital plate 
bearing 40-80 hairs. On Picea ...... piceae (p. 150) 

- Eighth abdominal tergite bearing 7-77 hairs but if with less than 19 (acuti- 

rostris, boerneri, cupressi, stroyani, pint, escherichi, fresai) then the fourth 
rostral segment is 130-2800 long, and if more than 2500 long (stroyani) then 
the fifth abdominal tergite bears 40-90 hairs between the siphunculi and the 
longest hair on the third abdominal tergite is 60-1500 long ... 8 

8 (7) Fifth abdominal tergite bearing only 3-8 hairs between the siphunculi. Meso- 

sternal tubercle well developed. 

Longest hair on the third abdominal tergite 12-600 long. Third antennal 
segment 400-7000 long, 6-5-15 times as long as the longest hair, 25-950, 
borne on it. Base of the sixth antennal segment 100-1800 long and bearing 
8-16 hairs. Second antennal segment bearing 5-13 hairs. Eighth abdominal 
tergite bearing 7-23 hairs. Second segment of hind tarsus 210-3200 long, 
2-0-2 -6 times as long as the first segment of the hind tarsus, which is 90-1500 
long, and 2 -7-3 -6 times as long as its own basal diameter. Second segment of 
hind tarsus 1-1-1-7 times as long as the fourth rostral segment, which is 
140-2500 long and bears 6-10 accessory hairs. Hind tibia 15-3-3 mm 
long and 23-44 times as long as thelongest hair, 40-120(1, borne on it. OnPinus 9 

Fifth abdominal tergite bearing 16—95 hairs between the siphunculi. 

Mesosternal tubercle absent ....... n 

9 (8) Processus terminalis bearing 5-7 sub-apical setae. Second antennal segment 

bearing 9-13 hairs. Eighth abdominal tergite bearing 14-23 hairs, the 
longest of which are 90-1 200 long. Hairs on the third abdominal tergite 
12-200 long. Body 3-4-4-3 mm long. Hind tibiae 2-6-3-3 mm long- 
Third antennal segment 9-5-15 times as long as the longest hair borne on 
it, which is 40-700 long. 

Fourth rostral segment 210-2300 long, 2-0-2-4 times as long as the fifth 
rostral segment and bearing 8-1 1 accessory hairs. On Pinns sylvestris. 

escherichi (p. 139) 

- Processus terminalis bearing only 4 sub-apical setae. Second antennal segment 

bearing 5-10 hairs. Eighth abdominal tergite bearing 7-18 hairs. Longest 
hair on third abdominal tergite 15-33 or 45 - 6°( x long. Either smaller, body 
2 -5-3-4 mm long, hind tibiae 1 -6-2-5 mm l° n g an d bearing hairs 40—950 long, 
third antennal segment 77-15 times as long as the longest hair, 25-800, 
borne on it, or body 3-5-4-1 mm long, hind tibia 2-7-3-2 mm long and bearing 
hairs 90-1200 long, and third antennal segment 6-5-9-5 times as long as the 
longest hair, 70-950 long, borne on it . . . . . . . 10 

10 (9) Body 3-5-4-1 mm long. Hind tibia 2-7-3-2 mm long. Longest hair on third 

abdominal tergite 45-650. long. Third antennal segment (Text-fig. 6) 550- 
6500- long, 6-5-9-5 times as long as the longest hair borne on it. Fourth 
rostral segment 210-2500 long (Text-figs 9 and 10). On Pinus nigra. 

acutirostris (p. 128) 
Body 2-5-3-4 mm l° n g- Hind tibiae 1-6-2 -5 mm long. Longest hair on third 
abdominal tergite 15-350 long. Third antennal segment 400-6000 long and 
7-5-15 times as long as the longest hair, 25-800, borne on it. Fourth rostral 
segment 140-2200 long. On Pinus sylvestris .... pini (p. 160) 

11 (8) Third abdominal tergite bearing hairs up to 12-350 long. Processus 

terminalis 14-25% of the total length of the sixth antennal segment. Fourth 
rostral segment 150-2100 long. Hind tibiae 2-3 mm long and 30-35 times 
as long as the longest hair, 50-700, borne on them. Second segment of hind 
tarsus 360-4600 long, 2-1-2-7 times as long as the first segment of the hind 



REVIEW OF CINARA IN BRITAIN 113 

tarsus, which is 150-2000 long. Fifth abdominal tergite bearing 16-36 
hairs between the siphunculi (Text-fig. 17). Primary rhinaria without 
chitinised rims. Third antennal segment (Text-fig. 18) without rhinaria. 
Hairs on the third antennal segment 20-700 long and on the eighth 
abdominal tergite 60-1000 long. On Larix . . . boerneri (p. 129) 

Hairs on third abdominal tergite up to 45-2700 long, if less than 600 then the 
processus terminalis is about 25% of the total length of the sixth antennal 
segment, the fourth rostral segment is 290-3300 long, the hind tibiae are 
2-3 mm long and 15-30 times as long as the longest hair, 1 10-1500, borne on 
them (schimitscheki) . If the longest hair on the third abdominal tergite is 
60-800 long, then either schimitscheki as previously or stroyani, in which case 
the fifth abdominal tergite bears 40-90 hairs between the siphunculi, the 
primary rhinaria have chitinized rims, the fourth rostral segment is 210-2800 
long, the third antennal segment usually bears a rhinarium and bears hairs 
up to 55-1 360 long and the eighth abdominal tergite bears hairs up to 
85-1600 long. On Pinus nigra, Abies, Picea, Cedrus and Cupressaceae . 12 

(11) Body 4-6-8 mm long. Hind tibiae 3-3-5-5 mm long and 12-20 (rarely to 30) 

times as long as the longest hair, rarely 150-1800 but usually 260-3300., 
borne on them. Third antennal segment (Text-fig. 1) 0-75-1-1 mm long 
and 2-5-4-5 times as long as the longest hair, 190-3000 borne on it. Base of 
the sixth antennal segment (Text-fig. 2) 230-2900 long and bearing 9-14 
hairs. Processus terminalis (Text-fig. 3) 44-900 long and bearing 3 or 4 
sub-apical setae. Fourth rostral segment (Text-fig. 4) 270-3500 long, 
2-1-2-6 times as long as the fifth rostral segment (120-1500), and bearing 
7-13 accessory hairs. Second segment of hind tarsus (Text-fig. 5) 380-5000 
long and 2-5-3 times as long as the first segment of the hind tarsus, which is 
130-1800 long. Third antennal segment without but fourth segment with 
1-4 rhinaria. Primary rhinarium of the fifth antennal segment without a 
chitinized rim. Fifth abdominal tergite bearing 70-90 hairs between the 
siphunculi, eighth abdominal tergite bearing 25-45 hairs up to 190-3600 
long. Siphuncular cones 460-9900 in diameter. On Abies spp. and some- 
times Cedrus ........ abieticola (p. 123) 

Body 17-50 mm long. Hind tibiae 09-30 mm long. Third antennal 
segment 250-9500 long. Base of the sixth antennal segment 110-2300 long. 
If body more than 4-5 mm long (bogdanovi, pilicornis, schimitscheki and 
fundatrices of stroyani) then the second segment of the hind tarsus is 1 -6-2-5 
times as long as the first segment of the hind tarsus [bogdanovi & schimitscheki) 
or 3-0-4-1 times as long as the first segment of the hind tarsus (pilicornis) or 
2-6—3-1 times as long as the first segment of the hind tarsus but in this case 
the siphuncular cones are 210-3700 in diameter, the third antennal segment 
is only 440-5200 long and is 4-5-7 times as long as the longest hair, 70-1000, 
borne on it, the base of the sixth antennal segment is only 130-1500 long, the 
processus terminalis is only 19-320 long, the fourth rostral segment is only 
220-2500 long, the fifth rostral segment is 90-1100 long, the hind tibiae are 
only 1 -7-2-1 mm long and bear hairs up to 100-1250 long and the longest hair 
on the third abdominal tergite is 80-1100 long and on the eighth abdominal 
tergite is 125-1550 long. On Pinus nigra, Picea spp. and Cupressaceae . 13 

(12) Second segment of hind tarsus 320-3800 long and 1 -6-i -9 times as long as the 

first tarsal segment which is 170-2200 long and 3-5-4-1 times as long as its 
own basal diameter. Second segment of hind tarsus 0-9-1 -2 times as long as 
the fourth rostral segment (Text-fig. 36) which is 290-3400 long and bears 6 
accessory hairs. Second antennal segment bearing 9-13 hairs and base of the 
sixth antennal segment bearing 6—9 hairs. 

Body 3-2-5-2 mm long and densely covered with small dark scleroites 



ii 4 v - F - EASTOP 

(Text-fig. 35) with a diameter only a little greater than the hair bases they 
encircle. Third antennal segment 6oo-8oo;i long and 5-5-12 times as long 
as the longest hair, 70-1 20[i, borne on it. Hind tibiae 2-3 mm long and 
15-30 times as long as the longest hairs, 110-150(1., borne on them. Longest 
hairs on the third abdominal tergite 45-100(1. Fifth abdominal tergite 
(Text-fig. 34) bearing 25-60 hairs between the siphunculi, eighth abdominal 
tergite bearing 25-65 hairs up to 95-1 300. long. Third and fourth antennal 
segments without rhinaria. Processus terminalis 55-70(1 long and about 
25% of the total length, 210-270(1, of the sixth antennal segment. On Pinus 
nigra .......... schimitscheki (p. 164) 

— Second segment of hind tarsus 200-520(1 long and 1 -9-4-1 times as long as the 

first segment of the hind tarsus, which is 6o-8ou. long and 1-5-3-8 times as 
long as its own basal diameter. If the fourth rostral segment is more than 
28o[i. long (bogdanovi) then it bears 9-13 accessory hairs and the second 
antennal segment and the base of the sixth antennal segment each bear 
11-26 hairs. On Picea spp. and Cupressaceae . ..... 14 

14 (13) Hind tibiae 1-3-2-7 mm long and 13-29 times as long as the longest hair, 

70-165(1, borne on it. Third antennal segment 340—740(1 long and 3-2-7-5 
times as long as the longest hair, 55-145(1, borne on it. Fourth rostral 
segment 210-320(1 long and bearing 6-13 accessory hairs. Primary rhinaria 
with a chitinised rim. On Picea . . . . . ... 15 

— Hind tibiae 0-9-2-2 mm long and 4-12 times as long as the longest hair, 

140—280(1, borne on it. Third antennal segment 240-580(1 long and 1-5-4-5 
times as long as the longest hairs borne on it, which are sometimes only 
8o-ioo[i but are usually 100-250(1 long; the ratio is more than 2-9 only in 
pilicornis. Fourth rostral segment 120-250(1 long and bearing 2-8 accessory 
hairs. On Picea spp. and Cupressaceae ....... 16 

15 (14) Sub-genital plate bearing 18-33 hairs mostly placed laterally. Third antennal 

segment often without rhinaria but sometimes 1 or 2 are present. Second 
segment of hind tarsus 250-380(1 long and 0-9-1 -4 times as long as the fourth 
rostral segment, 240-320(1, which bears 9-13 accessory hairs bogdanovi (p. 132) 

— Sub-genital plate bearing 33-65 hairs. Third antennal segment usually bear- 

ing 1-3 rhinaria but sometimes these are absent. Second segment of hind 
tarsus 310-47051 long and 1 -2-1 -9 times as long as the fourth rostral segment, 
which is 210-280(1 long and bears 6-9 accessory hairs . stroyani (p. 165) 

16 (14) Primary rhinaria with chitinized rims. Processus terminalis usually bearing 

4 but sometimes only 3 sub-apical setae. Eighth abdominal tergite bearing 
20-77 hairs up to 120-200(1 long. Third antennal segment 250-530(1 long, 
2-0-4-5 times as long as the longest hair, 80-170(1, borne on it. On Picea . 17 

— Primary rhinaria without chitinized rims. Processus terminalis usually bearing 

only 3 but sometimes 2 or 4, sub-apical setae. Eighth abdominal tergite 
bearing 16-31 hairs up to 120-260(1 long. Third antennal segment 190-500(1 
long, 1-5-2-5 times as long as the longest hair, 110-250(1, borne on it. On 
Cupressaceae ........... 18 

17 (16) Second segment of hind tarsus 250-330(1 long and o-6-i-o as long as the 

diameter of the siphuncular cones, 330-530(1. Third antennal segment 
280-450(1 long, 0-8-1-2 times as long as the diameter of the siphuncular 
cones and 2-0-2-8 times as long as the longest hairs, 140-170(1, borne on the 
third antennal segment ........ costata (p. 134) 

— Second segment of hind tarsus (Text-fig. 29) 300-520(1 long, 1-2-2-5 times as 

long as the siphuncular diameter of 130-390(1. Third antennal segment 
(Text-fig. 30) 250-530(1 long, 1-2-2-2 times as long as the siphuncular 
diameter and 2-3-4-5 times as long as the longest hairs, 80-150(1, borne on 
the third antennal segment ...... pilicornis (p. 152) 



REVIEW OF CINARA IN BRITAIN 115 

(16) Hind tibiae dark only at the apex, the proximal three quarters pale. Processus 
terminalis 11-20% (exceptionally to 28%) of the total length of the sixth 
antennal segment. Base of the sixth antennal segment 1 10-1700 long and 
bearing 8-14 hairs extending over most of its length. 

Longest hairs on the third antennal segment 1 10-1700 long, on third 
abdominal tergite 100-1800 and on the eighth abdominal tergite 120-1900 
long. Second segment of hind tarsus 200-2800 long and 1 -4-1 -7 times as 
long as the fourth rostral segment, which is 140-1800 long and bears 5-8 
accessory hairs. Yellow-brown or sometimes darker aphids, on the under- 
sides of the branches, often near the trunk of Callitris, Chamaecyparis, 
Libocedrus and Thuja orientalis ..... tujafilina (p. 166) 

Hind tibiae dark at both base and apex at least. Base of the sixth antennal 

segment 130-2300 long and 2-5-5 times as long as the processus terminalis 19 

(18) Hind tibiae completely black or dark brown. Third antennal segment 240- 

4100 long, 0-5-1-1 (but rarely exceeding 0-9) times as long as the diameter 
of the siphuncular cone, 270-5900. 

Third antennal segment usually shorter than the fourth and fifth segments 
together. Fourth rostral segment 120-1700 long and bearing 3-5 accessory 
hairs. Processus terminalis 40-700. long, base of the sixth antennal segment 
150-2300 long and bearing 3-5 accessory hairs. Sub-genital plate bearing 
19-27 hairs. Second segment of hind tarsus 250-3500 long, 3-1-4-0 times 
as long as the first segment of the hind tarsus, 70-1000, and 1 -8-2-4 times as 
long as the fourth rostral segment, which is 1-4-2-1 times as long as the first 
segment of the hind tarsus. On Juniperus communis . juniperi (p. 141) 

Hind tibiae with a paler area from about the basal one-fifth to half its length. 
Third antennal segment usually longer, ratio 0-7-1-7, than the diameter of 
the siphuncular cone. On Cupressus spp., Thuja occidentalis and Juniperus 
spp., other than communis ......... 20 

(19) Base of the sixth antennal segment 130-1600 long and bearing 4-7 (usually 5 or 

6) hairs, which are confined to the basal half. Fourth rostral segment 
120-1700 long, bearing 2-4 accessory hairs, and 1-4-1-9 times as long as the 
first segment of the hind tarsus. Sub-genital plate bearing 22-30 hairs. 
Smaller, body 2-3-5 mm long, pale reddish brown aphids on Cupressus 
macrocarpus, Thuja occidentalis and Juniperus virginiana . cupressi (p. 136) 

Base of the sixth antennal segment i40-200o(-230o in alatiform specimens) 
long. and bearing 7-12 hairs extending over most of its length. Fourth 
rostral segment 160-2400 long, bearing 5-7 accessory hairs and 1-8-2-2 
times as long as the first segment of the hind tarsus. Sub-genital plate 
bearing 28-44 hairs. Larger, body 2-2-4-2 mm long, dark brown aphids 
living on the under sides of the small branches of Cupressus and Juniperus spp. 

fresai (p. 140) 

Key to the alatae viviparae of the British species of Cinara 

Fourth rostral segment 300-4000 long and bearing 24-34 hairs arranged in 4 
longitudinal rows. Processus terminalis bearing 5-8 subapical setae and 
second antennal segment bearing 21-32 hairs. Fifth abdominal tergite 
bearing 18-40 hairs between the siphunculi. Hairs on the 3rd abdominal 
tergite up to 30-600 long. 

Body length 4-5-5-5 mm long, third antennal segment 700-9000 long, 10-15 
times as long as the longer hair, 60-800, borne on it. Fifth antennal segment 
350-6000 long, 2-6-3-4 times as long as the base of the sixth antennal 
segment. Base of the sixth antennal segment bearing 15-22 hairs. Eighth 
abdominal tergite bearing 41-64 hairs, the more dorsal hairs being 35-600 



n6 V. F. EASTOP 

long and the more lateral hairs on the eighth tergite are 85-130(1 long. 
Scleroites absent or very small. On Larix . . . kochiana (p. 143) 

Fourth rostral segment 130-380(1 long, but bearing only 2-17 hairs which are 
arranged in only 2 longitudinal rows. Processus terminalis usually bearing 
only 3 or 4 subapical setae, if with 5-1 1, then second antennal segment bear- 
ing only 7-18 hairs, and if the fifth abdominal tergite bears more than 15 
hairs between the siphunculi, then the hairs on the third abdominal tergite 
are up to 130-200(1 long ......... 2 

2 (1) Large aphid, body length 4-5-6-5 mm. Hind tibia 4-0-6-4 .mm and 65-120 

times as long as the longest hair, 45-75(1, borne on it. Longest hair on the 
third abdominal tergite 35-80(0.. Fifth abdominal tergite bearing 6-14 hairs 
between the siphunculi. Third antennal segment 0-65-1-3 mm long, 2-0-2-7 
times as long as the second segment of the hind tarsi, which are 450—520(1 
long; 2-7-3-3 times as long as the fourth rostral segment, and 17-26 times as 
long as the longest hair, 40— 65 jx, borne on the third antennal segment. 
Processus terminalis bearing 7-1 1 subapical setae. 

Fourth rostral segment, 300-370(1 long and bearing 10—17 accessory hairs. 
Fifth antennal segment 460-580(1 long, 1-8-2-3 times as long as the sixth 
antennal segment. Base of the sixth antennal segment 0-5-0-7 times as long 
as the fourth rostral segment and bearing 20—20 accessory hairs. Hind 
tarsus 1, 2-3-3-6 times as long as its basal diameter. Scleroites absent or 
very small. On Picea sp. ....... piceae (p. 150) 

- Body length 2-0-7-5 mm, but if more than 4-3 mm, then the longest hair on the 

third abdominal tergite is 80-320(1 long and if only 80-1 20(1 (laricis), then the 
dorsal abdominal hairs usually arise from conspicuous scleroites and the third 
antennal segment is 500-750(1 long, but only 1 -3-1 -9 times as long as the 
second segment of the hind tarsus and 7-12 times as long as the longest hair, 
50-80(1, borne on the third antennal segment. Fifth abdominal tergite 
bearing fewer than 8 or more than 15 hairs between the siphunculi. Pro- 
cessus terminalis usually bearing only 3 or 4 subapical setae but if with 6 or 7, 
then body 2 -8-4-4 mm l° n g. fifth antennal segment 280-360(1 long, hind tibiae 
2 -0-3 -3 mm long and 30-40 times as long as the longest hair, 75-95(1, borne 
on it (escherichi) or 14-17 times as long as the longest hair, 140—160(1, borne 
on it (brauni) ........... 3 

3 (2) Fifth abdominal tergite bearing 4-7 hairs between the siphunculi. Third 

antennal segment 400-700(1 long, 1-8-2-6 times as long as the second 
segment of the hind tarsus, 220—350(1; and 5-5-12 times as long as the longest 
hair, 50-100(1, borne on it. Scleroites absent. Body length 2-2-4-0 mm. 
On Pinus ............ 4 

- Fifth abdominal tergite bearing 16—70 hairs between the siphunculi. Third 

antennal segment 0-8-2-2 times as long as the second segment of the hind 
tarsus, if more than 1 -7 times then either the dorsal abdominal hairs arise 
from evident scleroites (laricis and schimitscheki) or the longest hairs on the 
third antennal segment are 140—320(1 long (bogdanowi and abieticola) . . 6 

4 (3) Fourth rostral segment 140-180(1 long, fifth rostral segment 70—95(1 long. 

Body length 2-2-4-0 mm. Hind tibiae 1-8-2-9 mm l° n g. !4- 2 5 times as 
long as the longest hair, 75-140(1, borne on them. Third antennal segment 
5-5-9-0 times as long as the longest hair borne on it. Longest hair on 3rd 
abdominal tergite 15-75(1, on 8th abdominal tergite 90-150(1. Second 
antennal segment bearing 6—9 hairs, base of sixth antennal segment bearing 
8-14 hairs. Fourth rostral segment bearing 6-10 hairs, subgenital plate 
bearing 20—42 hairs. Eighth abdominal tergite bearing 9-19 hairs. On 
Pinus sylvestris ......... pini (p. 160) 

- Fourth rostral segment 200-24051 long, fifth rostral segment 95-140(1 long . 5 



REVIEW OF CINARA IN BRITAIN 117 

Processus terminalis bearing 4 subapical setae. Second segment of hind 
tarsus 250-310^ long, 1 -1-1-4 times as long as the fourth rostral segment, 
which is 1 -6-i -8 times as long as the fifth rostral segment, 120-140^, 1-7-2-1 
times as long as the first segment of the hind tarsus, and bears 6-8 accessory 
hairs. Ultimate rostral segment (segments 4 + 5) 1-1-1-4 times as long as 
the second segment of the hind tarsus. Third antennal segment 2-6—3-0 
times as long as the fourth rostral segment, 6-8-5 times as long as the longest 
borne on it and bearing 1-6, usually 3-5 rhinaria. Hind tibia 2-5-3-2 mm 
long, 20-29 times as long as the longest hair, 1 io-i30[x, borne on it. Longest 
hair on the third abdominal tergite 65-1 oojz, on the 8th tergite 120-160^. 
Second antennal segment bearing 6-8 hairs, base of the sixth antennal 
segment bearing 9-13 hairs, subgenital plate bearing 24-38 hairs, and eighth 
abdominal tergite bearing 10-17 hairs. On Pinns nigra . acutirostris (p. 128) 

Processus terminalis bearing 6 or 7 subapical setae. Second segment of hind 
tarsus 300-36010. long, 1 -4-1 -7 times as long as the fourth rostral segment, 
which is 2-0-2-2 times as long as the fifth rostral segment, 95-1 iopL, 1-3-1-6 
times as long as the first segment of the hind tarsus and bearing 8-1 1 
accessory hairs. Ultimate rostral segment (segments 4 + 5) 0-9-1-1 times as 
long as the second segment of the hind tarsus. Third antennal segment 
3-0-3-3 times as long as the fourth rostral segment, 8-5-11-5 times as long as 
the longest hair borne on it and bearing 4-10, usually 5-7, rhinaria. Hind 
tibia 2-9-3-2 mm long, 32-38 times as long as the longest hair, 75-95^, borne 
on it. Longest hairs on the third abdominal tergite 30-55^, on the eighth 
tergite i20-i6o(x. Second antennal segment bearing 7-13 hairs, base of 
sixth antennal segment bearing 12-16 hairs, subgenital plate bearing 39-58 
hairs, eighth abdominal tergite bearing 17-22 hairs. On Pinus sylvestris. 

escherichi (p. 139) 

Base of the sixth antennal segment 250-320^ long and bearing 10-14 hairs. 
Hind tibiae 4-3-6-7 mm long and 14-24 times as long as the longest hair, 220- 
28o(i, borne on them. Large aphid, body length 5 -0-7 -5 mm. Third antennal 
segment 0-8-1-2 mm long, 2-9-4-5 times as long as the longest hair, 220-320^, 
borne on it. Longest hair on third abdominal tergite 160-320^, on eighth 
abdominal tergite 250-320^1. Primary rhinaria without chitinized rims. 
Fourth rostral segment 290-370^ long and bearing 8-13 accessory hairs. 
Second segment of hind tarsus 420-570^ long, 2-6-3-5 times as long as the 
first tarsal segment which is 140-200^ long and 2-2-2-9 times as long as its 
own basal width. Second antennal segment bearing 18-25 hairs. Third 
antennal segment bearing 7-15, usually 8 or 9, rhinaria. Fourth antennal 
segment bearing 2-5, usually 3 or 4, rhinaria. Scleroites very small. On 
Abies and Cedrus ........ abieticola (p. 123) 

Base of the sixth antennal segment 100-240^ long. Hind tibia 1 -3-4-1 mm long. 
Body length 2-0-5-3 mm, but if more than 45 mm, then primary rhinaria 
with a chitinized rim, longest hair on the third antennal segment less than 
20o\i long and on eighth abdominal tergite less than 250(0. long. If the body 
is more than 4-5 mm long then either it is 2-0-2-3 times as long as the hind 
tibiae, 1-9-2-2 mm (pectinatae) , or the hind tibiae are 2-2-3-2 mm long 
{laricis and pinea) and bear hairs only up to I20-I50[x long (laricis) or the 
first tarsal segment is 210—330(7. long and 4-3-5-7 times as long as its own 
basal width (pinea) or the hind tibiae are 2-4-3-5 mm l° n g an d the second 
tarsal segment 310-420(0. long and 2-2-2-7 times as long as the first tarsal 
segment (bogdanowi) .......... 7 

Rostral segments 4 + 5 together 1-3-1-5 times as long as the second segment of 
the hind tarsus, which is 0-9-1 -2 times as long as the fourth rostral segment 
alone. Fourth rostral segment 300-370(0 long and bearing 6 or 7 accessory 



u8 V. F. EASTOP 

hairs. Second segment of hind tarsus 1-7-2-0 times as long as the first 
segment of the hind tarsus, 160-2000- Second antennal segment bearing 
7-13 hairs, base of the sixth antennal segment bearing 6—9 hairs, processus 
terminalis with 4 subapical setae and sub-genital plate bearing 50-70 hairs. 
Fifth abdominal tergite bearing 45-80 hairs between the siphunculi, eighth 
abdominal tergite bearing 27-60 hairs. Dorsal abdominal hairs arising from 
small dark scleroites 12-200 in diameter, about twice the diameter of the hair 
bases. Third antennal segment 1 -8-2-1 times as long as the second segment 
of the hind tarsus without or bearing only one rhinarium. Longest hair on 
eighth abdominal tergite 1 10-1500 long, 0-8-1-2 times as long as the longest 
hair, 120-1500 on the third antennal segment. Longest hair on the hind 
tibiae 150-1700 and on the third abdominal segment 70-1100- Fifth 
antennal segment 1 -2-1 -5 times as long as the sixth segment and 0-4-0-6 as 
long as the fourth rostral segment which is 1 -4-2 o times as long as the first 
segment of the hind tarsus. On Pinns nigra . . . schimitscheki (p. 164) 

— Rostral segments 4 + 5 together 0-5-1 -3 times as long as the second segment of 

the hind tarsus which is rarely less than 1 3 times as long as the fourth 
rostral segment alone. Fourth rostral segment 130-3100. long but if more 
than 2600 then the longest hair on the third abdominal tergite is 120-2400 
long and either the fourth rostral segment bears 8-15 accessory hairs, the 
longest hairs on the third antennal segment are 140-1900, on the eighth 
abdominal tergite are 150-2200, scleroites are absent, the second antennal 
segments bear 14-24 hairs, the base of the sixth antennal segment bears 11-19 
hairs, the subgenital plate bears 22-40 hairs and the second segment of the 
hind tarsus is 2-2-2-7 times as long as the first segment of the hind tarsus, or 
if the fourth rostral segment bears 4-7 accessory hairs, then the third 
antennal segment is 1 -2-1 -7 times as long as the second segment of the hind 
tarsus, the longest hair on the eighth abdominal tergite is 1 -2-1 -9 times as 
long as the longest hair on the third antennal segment, the longest hairs on the 
hind tibiae are 190-2700 long and the longest of 17-27 hairs on the eighth 
abdominal tergite are 170-2500 long, the sixth antennal segment is 0-65-1-0 
times as long as the first segment of the hind tarsus, which is 210-3300 long 
and the dorsal abdominal hairs arise from large scleroites (pinea), or the 
processus terminalis bears 6 or 7 hairs, the base of the sixth antennal segment 
bears n-14 hairs, the fifth abdominal tergite bears 16-26 hairs between the 
siphunculi, the eighth abdominal tergite bears n-16 hairs, the third antennal 
segment bears hairs up to 80-1100 long and usually 2-4 rhinaria (brauni) . 8 

8 (7) A short-haired aphid. Third antennal segment 550-7500 long and 17-25 times 
as long as the longest hair, 20-450, borne on it. Hind tibia 2 -6—3-2 mm long, 
35-70 times as long as the longest hair, 50-800, borne on it. Longest hair on 
the 3rd abdominal tergite only 20-400 long, on the eighth abdominal tergite 
70-1000. Fifth abdominal tergite bearing 16-25 hairs between the 
siphunculi. Second segment of hind tarsus 440-5700 long, 2-4-2-8 times as 
long as the fourth rostral segment, and 1 -o-i -8 times as long as the diameter 
of the siphuncular cone. 

Processus terminalis 12-20% of the total length of the sixth antennal 
segment. Base of the sixth antennal segment 150-1900 long, 0-9-1-2 times 
as long as the fourth rostral segment and bearing 7-9 hairs. Fourth rostral 
segment bearing 4-8 accessory hairs. Dorsal abdominal hairs arising from 
small scleroites at the hair bases. On Larix .... boerneri (p. 129) 

- Longer-haired aphids. Third antennal segment 300-9000 long and 1-5-12 

times as long as the longest hair, 50-2500, borne on it. Hind tibia 1-4- 
3-5 mm long, 5-5-23 times as long as the longest hair, 95-3500, borne on it. 
Longest hairs on the third abdominal tergite 70-2300 long, on the eighth 



REVIEW OF CINARA IN BRITAIN 119 

abdominal tergite 100-260JX long, if less than 130^ the fifth abdominal 
tergite usually bearing 24-54 hairs between the siphunculi. Second segment 
of the hind tarsus less than 2 -4 times as long as the fourth rostral segment or, 
if longer, then the body and appendages usually bear long fine hairs, those on 
the antennae being 85-230^, on the hind tibiae 180-350^, on the third 
abdominal tergite 100-230(1. and on the eighth abdominal tergite 120-260U. 9 

Dorsal abdominal hairs arising from conspicuous dark scleroites (Text-fig. 32), 
45-1 20(j. in diameter, the larger scleroites being 3-6 times the diameter of the 
hair bases borne on them. Third antennal segment is 2-2-3-0 times as long 
as the fourth rostral segment. Longest hair on third antennal segment 
I20-I70(i, on third abdominal tergite 150-240(0. long. The fifth rostral 
segment is 210-330U. long. Fifth antennal segment 1-1-1-7 times as long as 
the sixth antennal segment. Base of the sixth antennal segment 150-230^, 
0-65-0-95 times as long as the fourth rostral segment. Hind tibiae 2-2- 
3-2 mm long, 9-16 times as long as the longest hair, 190-270^, borne on them. 
Body length 3-2-5-3 mm. Third antennal segment 500-800U, long, 3-5- 
6-o times as long as the longest hair borne on it and 1 -2-1 -7 times as long as 
the second segment of the hind tarsus. Processus terminalis 40-85(1. long and 
bearing 4 sub-apical setae. Fourth rostral segment 210-270(1. long, bearing 
4-6 accessory hairs and 1-6-2-1 times as long as the fifth rostral segment. 
Second segment of hind tarsus 1 -4-2 -o times as long as the first segment of 
the hind tarsus which is 4-3-5-7 times as long as its basal width. On Pinus 
sylvestris .......... pinea (p. 156) 

Dorsal abdominal scleroites small or absent or, if large and conspicuous, then the 
third antennal segment is 3-0-4-3 times as long as the fourth rostral segment, 
the longest hair borne on the third antennal segment is 50-1 20fz long, and on 
the third abdominal tergite is 70-150(1. long and the fifth rostral segment is 
85-1 1 ou. long; and either the third antennal segment is 7-12 times as long as 
the longest hair borne on the third segment, the hind tibia is 2-2-3-0 mm 
long, 16-23 times as long as the longest hair borne on it, the base of the sixth 
antennal segment is ioo-i6oji. long, the processus terminalis is 25-45(1. long, 
the fourth rostral segment is 160-200(1. long and the fifth rostral segment is 
85-110}/ long (laricis), or the first segment of the hind tarsus is 130-180(1. long 
and 2-6-4-0 times as long as its own basal diameter, the longest hair on the 
hind tibia is 120-150U., the third antennal segment is 300-550(1 long and 
1 -8-2-1 times as long as the second segment of the hind tarsus and 1-6-2 -3 
times as long as the fourth rostral segment, which is 2-1-2-8 times as long as 
the fifth rostral segment; the fifth antennal segment is o-8-i 1 times as long 
as the sixth antennal segment, the base of the sixth antennal segment is 
i6o-200(x long and 0-45-0-6 times as long as the fourth rostral segment, 
whish is 190-250U. long and bears 7-8 accessory hairs (pectinatae) . . 10 

Processus terminalis bearing 6 or 7 sub-apical setae. Fourth rostral segment 
260-300U. long and bearing 4-7 accessory hairs. Second antennal segment 
bearing 8-12 hairs. Fifth abdominal tergite bearing 16-20 hairs between the 
siphunculi, eighth abdominal tergite bearing 12-16 hairs. 

Third antennal segment 1 -3-1 -7 times as long as the second segment of the 
hind tarsus, 6-5-8-0 times as long as the longest hair, 80-1 iO(i., borne on it 
and bearing 9-13 rhinaria. Longest hair on the eighth abdominal tergite 
130-200(1., 1-5-1-9 times as long as the longest hair on the third antennal 
segment. Fifth antennal segment 1 -o-i -2 times as long as the sixth antennal 
segment. First tarsal segment 4-5 times as long as its basal diameter. 
Scleroites absent. Body 2-8-3-7 mm l° n g- Hind tibiae 2-1-2-5 mm long, 
14-17 times as long as the longest hair, 140-160(1., borne on them. On Pinus 
nigra ........... brauni (p. 133) 



120 V. F. EASTOP 

— Processus terminalis bearing 3 or 4, or exceptionally 2 or 5 sub-apical setae. 

Fourth rostral segment usually only 130-240^ long but if 240-290^ long, 
then bearing 8-15 accessory hairs and the second antennal segment bears 
14-26 hairs, the fifth abdominal tergite bears 30-55 hairs between the 
siphunculi and the eighth abdominal tergite bears 24-42 hairs (bogdanowi) . 1 1 

11 (10) Fourth rostral segment 230-300^ long and bearing 8-15 accessory hairs. 

Third antennal segment 500-900^1 long, and 1-7-2-2 times as long as the 
second segment of the hind tarsus, which is 2-2-2-7 times as long as the first 
segment of the hind tarsus. Fourth antennal segment 240-440^ long, i-i— 
1-7 times as long as the sixth antennal segment, fifth antennal segment 
300-480^ long and 1 -5-1 -9 times as long as the sixth antennal segment. 

Body 3-2-5-0 mm long. Siphuncular cone 500-7000. in diameter. Hind 
tibiae 2-4-3-5 mm long. Processus terminalis bearing 4 subapical setae. 
On Picea ......... bogdanowi (p. 132) 

— Fourth rostral segment 130-250^ long and bearing 2-8 accessory hairs. If 

fourth rostral segment more than 2IO(jl long, then the third antennal segment 
is 300-660JJ. long, o-8-i-6 times as long as the second segment of the hind 
tarsus which is 2-8-4-4 times as long as the first segment of the hind tarsus, 
the fourth antennal segment is 120-240(1 long and 0-7-1-2 times as long as 
the sixth antennal segment, the fifth antennal segment is 160-280^ long and 
0-8-1-4 times as long as the sixth antennal segment .... 12 

12 (11) Third antennal segment 500-750^ long, 7-5-12-0 times as long as the longest 

hair, 5c—8o[i., borne on it. Dorsal abdominal hairs usually arising from dark 
scleroites, the largest of which have a diameter several times that of the 
bases of the hairs borne on them. Fifth antennal segment 1-7-2-7 times as 
long as the sixth antennal segment. Longest hair on the third abdominal 
tergite 70-1 iojj. long. Third antennal segment 1-3-1-9 times as long as the 
second segment of the hind tarsus, which is 2-1-2-7 times as long as the first 
segment of the hind tarsus. 

Body length 3-1-4-8 mm. Hind tibiae 2-2-3-0 mm long, 16-23 times as 
long as the longest hair, 120-1500., borne on them. Longest hair on the 
eighth abdominal tergite 1 -7-3 -o times as long as the longest hair on the third 
antennal segment. On Larix ...... laricis (p. 146) 

— Third antennal segment 1-5-7-0 times as long as the longest hair borne on it. 

Dorsal abdominal hairs without or with only very small scleroites at their 
bases, or if the scleroites are more than twice the diameter of the hair bases 
(pectinatae) then the fifth antennal segment is o-8-i-i times as long as the 
sixth antennal segment and the second segment of the hind tarsus is 1-7-2-1 
times as long as the first segment of the hind tarsus. Fifth antennal segment 
usually 0-7-1-4 times as long as the sixth segment, if 1-5-2-0 times as long 
(piniliabitans) the longest hair on the third abdominal tergite is i20-i8o[x 
long, on the third antennal segment is 120-1601J. long, and the third antennal 
segment is 2-0-2-5 times as long as the second segment of the hind tarsus. 
Hind tibia 5-5-16 (exceptionally — 18) times as long as the longest hair borne 
on it. Longest hair on the eighth abdominal tergite 0-8-2 -o times as long as 
the longest hair on the third antennal segment, except in pectinatae . . 13 

13 (12) Many dorsal abdominal hairs arising from scleroites, the largest of which have 

a diameter three times that of the hair bases. Dorsal length of first segment 
of hind tarsus 150-180^, ventral length 190-270^ and 3-9-5-6 times as long 
as its proximal diameter. Body 3 -2-5 -o mm long and 2 -0-2-3 times as long as 
the hind tibiae, which are 1-4-2-2 mm long. Longest hairs on the eighth 
abdominal tergite 170-210^ long and 1-6—2-8 times as long as the longest 
hairs on the third antennal segment, which are 75-120^ long. Fourth 
rostral segment 190-250^ long, 2-1-2-8 times as long as the fifth rostral 



REVIEW OF CINARA IN BRITAIN 121 

segment and o-8-i-i times as long as the first segment of the hind tarsus. 
Fifth antennal segment o-8-i 1 times as long as the sixth. Second antennal 
segment .bearing 6-8 hairs, base of the sixth antennal segment bearing 5-7 
hairs. Green aphid on A bies . ..... pectinatae (p. 148) 

Dorsal abdominal scleroites absent or, if present, not exceeding twice the 
diameter of the hair bases. First segment of hind tarsus 15-80(0. long 
dorsally, 80-1 6ou long ventrally, and 2-1-3-6 (exceptionally 4-0) times as 
long as its proximal diameter. Body 2-0-4-5 mm long, 1-2-2-2 times as 
long as the hind tibiae, only exceeding 2 -o in pilicornis. Longest hair on the 
eighth abdominal tergite 100-260(0. long, 1-0-1-7 (rarely 2-0) times as long as the 
longest hair, 70-250(0., on the third antennal segment. Fourth rostral segment 
1-5-2-2 times as long as the fifth rostral segment and usually 1-2-2-4 as l° n g 
as the first segment of the hind tarsus. If fourth rostral segment 1 0-1 3 
times as long as the first segment of the hind tarsus (pinihabitans), then fifth 
antennal segment 1-5-20 times as long as the sixth antennal segment. 
Second antennal segment bearing 5-20 hairs, base of the sixth antennal 
segment bearing 5-20 hairs. Brown, reddish or yellowish aphids on Pinus, 
Picea and Cupressaceae .......... 14 

(13) Third antennal segment 550-750(0 long, 3-8-5-2 times as long as the fourth 

rostral segment which is 130-170(0 long, bears 4 accessory hairs and is 1 -o-i -3 
times as long as the first segment of the hind tarsi. Third antennal segment 
2-0-7-5 times as long as the second segment of the hind tarsus, which is 
260-340(0. long and 2-0-2-3 times as long as the first segment of the hind 
tarsus. Fifth antennal segment 240-380(0. long and 1-5-2-0 times as long as 
the sixth antennal segment, which is 1 20-1600. long. Base of the sixth 
antennal segment bearing 7-1 1 hairs. Second antennal segment bearing 
6-8 hairs. 

Processus terminalis bearing 4 subapical setae. Eighth abdominal 
tergite bearing 11-22 hairs. On Pinus sylveslris . . pinihabitans (p. 162) 

Third antennal segment 300-6000 long, 1-8-3-6 times as long as the fourth 
rostral segment, which is 130-250(0 long and is 1-2-2-2 times as long as the 
first segment of the hind tarsus. Third antennal segment 0-9-1-6 times as 
long as the second segment of the hind tarsus, which is 240-540(1 long and 
2-6-4-4 times as long as the first segment of the hind tarsus. Fifth antennal 
segment 160-300(0. long and 0-7-1-4 times as long as the sixth antennal 
segment, which is 110-2400. long. Base of the sixth antennal segment 
bearing 5-20 hairs. Second antennal segment bearing 7-20 hairs. On 
Picea and Cupressaceae .......... 15 

(14) Membrane of the fore wing characteristically pigmented (Text-fig. 20), media 

once branched. Second segment of the hind tarsus 300-380(0 long, 1-3-1-7 
times as long as the fourth rostral segment which is 190-250(0 long. Primary 
rhinaria with a chitinized rim. Third antennal segment 2-0-3-5 times as 
long as the longest hair, 1 50-2100., borne on it, and 1-3-1-6 times as long as 
the second segment of the hind tarsus, which is 0-5-1 -o times as long as the 
siphuncular diameter. Hind tibia 2 -0-2 -6 mm long and 7 -5-10 times as long 
as the longest hair, 190-300(0, borne on it. Third antennal segment bearing 
only 1-3 rhinaria. Base of the sixth antennal segment bearing 12-20 hairs. 
Processus terminalis bearing 3 or 4 sub-apical setae. Processus terminalis 
only 6-18% of the total length of the sixth antennal segment. On Picea. 

costata (p. 134) 
Wing membrane uniformly transparent, media usually twice branched (Text- 
fig- 37)- Second segment of the hind tarsus 1-6-2-7 times as long as the 
fourth rostral segment, if less than 1 -8 then either the primary rhinaria 
without a chitinized rim (subgen. Cupressobium) or the longest hair on the 



122 V. F. EASTOP 

hind tibia only 95-180(1 long (stroyani). Primary rhinaria with or without a 
chitinized rim but if with, then the third antennal segment is 3-0-3-7 times 
as long as the longest hair, 70-160U., borne on it and 0-9-1 -3 times as long as 
the second segment of the hind tarsus and the base of the sixth antennal 
segment bears 6-17 but mostly 9-12 hairs. Processus terminalis 14-29% of 
the total length of the sixth antennal segment except in tnjafilina (8-15%), 
in which the fourth rostral segment is only 130-180(1 long. . . . 16 

16 (15) Longest hair on the third abdominal tergite 55-1 10(1 long. Hind tibiae 9-5-17 

times as long as the longest hair, 140-180(1, borne on the hind tibiae. Third 
antennal segment 3-5-7-0 times as long as the longest hair, 70-110(1, borne 
on it. Base of the sixth antennal segment bearing 9-17 hairs and o-6-o-8 
times as long as the fourth rostral segment. 

Processus terminalis bearing 4 sub-apical setae. On Picea stroyani (p. 165) 

- Longest hair on the third abdominal tergite 105-240(1 long. Hind tibiae 

5-5-11 but rarely more than 9-5 times as long as the longest hair, 140-360(1, 
borne on the hind tibia. If the longest hind tibial hair is less than 180(1 long 
(iujafilina), then the fourth rostral segment 130-180(1 long and the third 
antennal segment is 2-0-2-9 times as long as the longest hair, 120-200(1, 
borne on it, the longest hair on the third abdominal tergite is 120-210(1 long 
and the base of the sixth antennal segment is 0-9-1-3 times as long as the 
fourth rostral segment. Third antennal segment 1 -5-5 -o times as long as the 
longest hair which is rarely 80-1 ioji, but usually 110-250(1 long . . 17 

17 (16) Processus terminalis bearing 4 sub-apical setae. Primary rhinaria with a 

chitinized rim. Third antennal segment 30-3-5 times as long as the longest 
hair, 8o-i6o(i, borne on it. Base of sixth antennal segment o-6-o-g times as 
long as the fourth rostral segment which is 160-250, rarely less than 190(1, 
long. Longest hair on hind tibia 160-240(1. Second segment of hind tarsus 
350-420(1 and 1-1-2-3 times as long as the diameter of the siphuncular cone. 
On Picea ......... pilicornis (p. 152) 

- Processus terminalis bearing only 3 sub-apical setae. Primary rhinaria 

without a chitinized rim. Third antennal segment 1 -8-2 9 times as long as 
the longest hair on it. Base of the sixth antennal segment 0-9-1 -7 times as 
long as the fourth rostral segment, which is 130-210(1 long, but if more than 
i8o[i long, then the longest hair on the third antennal segment is 190-250(1 
long. Longest hair on hind tibia 140-350(1, if less than 230(1 (tujafilina), 
then the second segment of the hind tarsus is only 240-34051 long. On 
Cupressaceae (subgen. Cupressobium) ....... 18 

18 (17) Base of the sixth antennal segment bearing only 5 or 6 hairs. Fourth rostral 

segment 130-170(1 long and bearing 2-4 accessory hairs. Third antennal 
segment bearing 1-6 rhinaria, usually confined to the distal half of the 
segment. 

Fifth antennal segment 180-240(1 long. Processus terminalis 30-50(1 long, 
15-23% of the total length of the sixth antennal segment. On Cupressus, 
Juniperus and Thuja ........ cupressi (p. 136) 

- Base of the sixth antennal segment bearing 6-14 but rarely less than 8 hairs. 

Fourth rostral segment 130-210(1 long and bearing 4-8 accessory hairs. 
Third antennal segment bearing 3-1 1 rhinaria, usually extending onto the 
proximal half of the segment ......... 19 

19 (18) Tibiae pale to dusky, rarely dark brown, except for the distal apex which is 

dark. Processus terminalis 10-30(1 long, 8-16% of the total length of the 
sixth antennal segment. Third antennal segment (Text-fig. 38) 300-450(1 
long, 1 -o-i g but rarely less than 1 -3 times as long as the diameter of the 
siphuncular cone. Fifth antennal segment 160-200(1 long. Longest hair on 
eighth abdominal tergite 130-210(1 long. Second segment of hind tarsus 



REVIEW OF CINARA IN BRITAIN 123 

(Text-fig. 39) 240-330(0. long, 0-7-1-4, usually 10-13 times as long as the 
diameter of the siphuncular cone (Text-figs 40 & 41). On Callitris and 
Thuja .......... tujafllina (p. 166) 

Tibiae black or black at base and apex and dark brown in the middle. 
Processus terminalis 35-80(0. long, 14-29% of the total length of the sixth 
antennal segment. Third antennal segment 350-600(1. long, o-8-i-6 but 
rarely more than 1 3 times as long as the diameter of the siphuncular cone. 
Fifth antennal segment 180—300(0, rarely less than 200^ long. Longest hair 
on eighth abdominal tergite 190-260(0, rarely less than 210(0 long. Second 
segment of hind tarsus 280-400(0 long, 0-6-1-2, usually 0-7-1-0 times as long 
as the diameter of the siphuncular cone. On Cupressus and Juniperus . 20 

20 (19) Second segment of hind tarsus 1-6-2-1, usually 17-20 times as long as the 
fourth rostral segment, which bears 5-8 accessory hairs. Second antennal 
segment bearing 10-14 hairs. Fifth abdominal tergite bearing 60 or more 
hairs between the siphunculi. Third antennal segment 1-2-1-6, rarely less 
than 1 -3 times as long as the second segment of the hind tarsus. Base of the 
sixth antennal segment 0-8-1-3, usually 1-0-1-3 times as long as the fourth 
rostral segment. Longest hair on the eighth abdominal tergite 0-9-1-2 
times as long as the longest hair on the third antennal segment. Radial 
sector of fore wing often not reaching wing apex. On Cupressus and 
Juniperus .......... fresai (p. 140) 

Second segment of hind tarsus 2-0-2-8, usually 2-4-2-6 times as long as the 
fourth rostral segment, which bears 4 accessory hairs and is 1 -4-1 -6 times as 
long as the first segment of the hind tarsus. Second antennal segment 
bearing 7-1 1 hairs. Fifth abdominal tergite bearing 25-45 hairs between 
the siphunculi. Third antennal segment 1 -o-i -3 times as long as the second 
segment of the hind tarsus. Base of the sixth antennal segment 1-1-1-7, 
usually 1 4-1 -6 times as long as the fourth rostral segment. Longest hair on 
the eighth abdominal tergite 1 -2-1 4 times as long as the longest hair on the 
third antennal segment. Radial sector of fore wing reaching wing-apex. 
On Juniperus ......... juniperi (p. 141) 

The British Species of CINARA 

Cinara abieticola (Cholodkovsky) 

(Text-figs 1-5) 

Aphis borealis Curtis, 1828 : 201. [Types unknown. 'Floating floes of ice in the Polar Sea . . . 

as far north as 82 .'] 
Lachnus confinis Koch, 1856 : 245 [No types. Collection data not given, presumably 

Germany.] 
[Lachnus grossus (Kaltenbach) Cholodkovsky, 1896a : 145; 1898a : 656-657; 1902 : 6; 

Mordwilko, 1929 : 27; Gillette & Palmer, 1930 : 544, 551. Misidentifications.] 
Lachnus abieticola Cholodkovsky, 1899 : 470-471. [Types unknown. Western Siberia: 

Tomsk, Summer 1898, apterae viviparae (W. Plotnikov)]; 1902 : 6. 
Lachniella cilicica Del Guercio, 1909 : 287, 297-301. [Types unknown. Italy: Castelfalfi, 

May, 1893, aptera vivipara (Biondi)]; Jackson, 1919 : 164. 
Lachniella cilicica v. cecconii Del Guercio, 1909 : 297. [As for cilicica s. str.] 
Lachnus vanduzei Swain, 1919 : 50-51. [Types, University of California. California; 

spruce, September & November, apterae & alatae viviparae, 1914 (E. P. Van Duzee & 

E. 0. Essig).] 
[Dilachnus piceae (Walker) Swain, 1921 : 225-227, nee Panzer, 1801.] 
Panimerus cilicica (Del Guercio) Theobald, 1929 : 142-145. 
Panimerus vanduzei (Swain) Theobald, 1929 : 154-156, partim, the Irish specimens. 



124 



V. F. EASTOP 




Figs 1-5. Cinara abieticola, aptera vivipara. 1, third antennal segment, x 87. 2, sixth 
antennal segment, x 87. 3, processus terminalis, x 450. 4, fourth and fifth rostral 
segments, x no. 5, hind tarsus, x 87. 



REVIEW OF CINARA IN BRITAIN 125 

? Lachnus lasiocarpae Gillette & Palmer, 1930 : 543-544. [Types, USNM. Catalogue no. 

42082. 'Paratypes' at Fort Collins. Colorado: A bies lasiocarpa.] 
Dilachnus pubescens Wellenstein, 1930 : 743-747, 752. [Types unknown. Collection data 

not given, presumably Germany: Hann, alate males and oviparae.] 
? Cinara lasiocarpae (Gillette & Palmer) Gillette & Palmer, 1931 : 858; Palmer, 1952 : 34. 
Cinara cecconii (Del Guercio) Braun, 1938 : 478. 

[Cinara grossus (Kaltenbach) Blanchard, 1939 : 862-864. Misidentification.] 
Dinolachnus cecconii (Del Guercio) Borner, 1940 : 1. 
Neochmosis cilicica (Del Guercio) Kloet & Hincks, 1945 : 72. 
[Cinara grossa (Cholodkovsky nee Kaltenbach) Mordwilko, 1948 : 201-202.] 
Todolachnus abieticola (Cholodkovsky) Borner, 1952 : 44; Pasek, 1954 : 266-269; Heinze, 

1962 : 173; Gomez-Menor, 1962 : 390-393. 
Todolachnus confinis (Koch) Borner, 1952 : 44; Heinze, 1962 : 173. 

Cinara abieticola (Cholodkovsky) Hottes & Essig, 1954 : 95~97; Szelegiewicz, 1962 : 78-79. 
[Dinolachnus piceae (Panzer) Aizenberg, 1956 : 139; Pintera, 1966 : 281. Misidentincations.] 
Cinaria (Pitsaria) vanduzei (Swain) Aizenberg, 1956 : 139. 
Todolachnus abieticola subsp. bulgarica Pintera, 1959 : 71. [Holotype, Academy of Science, 

Prague. Bulgaria: Rila, Abies alba bulgarica, August, 1957, apterae viviparae {A. 

Pfeffer).] 
Cinara (Todolachnus) abieticola (Cholodkovsky) Heikinheimo, 1963 : 185; Canakcioglu, 

1966 : 140-141. 
[Cinara piceae (Panzer) Shaposhnikov, 1964 : 522. Misidentification.] 
Dinolachnus abieticola (Cholodkovsky) Hille Ris Lambers, 1966 : 124. 

Material examined. 

England: London, 'conifer', 15. ix. 1948, 1 aptera (A. Smith); 'near conifer', 
23. vi. 1969, 1 apt., 1 alata (B. J. English); Southgate, Cedrus deodora, i.vii.i955, 
4 al. (Parks Superintendent); Surrey, Chertsey, 'near cedar', March 1967, 8 apt. & 
1 alatoid nymph (Public Health Inspector); Essex, Havering, 'cedar', 12. v. 1967, 
1 apt., 1 al., 1 alatoid nymph (Health & Welfare Department); Worcs., Stourbridge, 
'crawling under door', 6. v. 1949; 2 apt., 1 immature (Ministry of Health); Devon, 
Germansweek, Beauworthy, 'fir', 4^.1948, 1 al. (E. W. Powell). Ireland: Avon- 
dale, Rathdrum, Pinus, 2.vi.i9i3, 1 apt., 2 al. (F. V. Theobald). Scotland: Moray, 
Rothes, Abies pectinata, 24-27. ix.1917, 1 apt. (D. J. Jackson); Elgin, Pitgaveny, 
Abies nobilis -\- A. grandis, 12. vi. 1923, 2 apt., 3 al., 4 alatoid nymphs (A. Stables); 
Roxburgh, Craik Forest, Abies alba, 6.iii.i964, 1 apt. (C. I. Carter); Argyll, Kilmum 
arboretum, Abies lasiocarpa, 9.vii.i965, 1 apt. (C.I.C.). 

Czechoslovakia: Kosice-Crmel, Abies alba, 3^.1948, 2 apt. (V. Pasek). Den- 
mark: Palagaard Forest, A. alba, 18. vi. 1964 (C. I. Carter). France: Vosges, 
Retournermer, 7-31.vii.1930, 2 al. (M. E. Mosely). Germany: Unteres Weldental, 
A. alba, 15.vi.1961, 1 apt. (H. Heinze). Spitzbergen: W. of N.E. Land, S. of Lee- 
cap, N. of Wahlenberg Bay, c. 1000 ft, 8.viii.i924, 1 al. (H. M. Clutterbuck) (other 
specimens from this series in the Hope Museum, Oxford). Turkey: Bolu-Aladag, 
1360 m., Abies bornmuelleri , 4.^.1964, 18 apt., 1 alatoid nymph (H. Canakcioglu). 
India: Mussourie, 6500 ft, under bark of Cedrus deodora, July 1920, 2 al., 2 alatoid 
nymphs (S. N. Chatter jee) ; Rhotang Pass, 13,800 ft, 4.VL1955, 3 al. (A. P. Kapaur). 
Pakistan: Murree, Abies pindrow, 2 adult and 2 immature oviparae, 25. xi. 1958 
(M. Ghani); Murree, in spider's web on fir, 2 alate males, 25.xi.1958 (M. Ghani). 



126 V. F. EASTOP 

U.S.A.: California, Berkeley, spruce, 2.ix.i9i4, i aptera [cotype of vanduzei]; on 
pavement, 3.iii.io,6i, 2 al. (0. W. Richards). Canada: Labrador, Mealy Mts, 
31.viii.1958, 2 oviparae (British School Boys' Exploration Society); New Brunswick, 
Acadia Stn, Abies balsamea, 8.vi.i96o, 2 apt. (M. E. MacGillivray). 

Host-plants. Specimens have been seen from Abies alba (= pectinata), 
A. balsamea, A. bornmuelleriana, A. grandis, A. lasiocarpa, A. nordmanniana, 
A . pindrow and Cedrus deodora. Cinara abieticola was described from Abies sibirica, 
(as Lachnus cilicica) from A. cilicica, and is also recorded from Abies procera 
(= nobilis) and A. religiosa. 

Distribution. Specimens have been seen from Czechoslovakia, Denmark, 
England, France, Germany, Ireland, Scotland, Spitzbergen, Turkey, India, Pakistan, 
California, Labrador and New Brunswick. Cinara abieticola was described from 
Western Siberia, as Lachnus cilicica from Italy, as bulgarica from Bulgaria and is 
also recorded from Austria, Finland, Norway (Stenseth & Bekke, 1968 : 237), 
Portugal (Ilharco, 1968a : 118), Spain, Sweden, Yugoslavia, U.S.S.R., Latvia 
(Rupais, 1961 : 16-17), Ukraine (Mamontova, 1963 : 21, as piceae), Argentina 
(Blanchard, 1939 : 862-864, as grossa) and the Eastern and Western coasts of 
North America (Hottes, i960 : 221). Borner (1952 : 44) lists the Italian synonym 
but says that it is not certain that Italian specimens really belong to the northern 
European species. 

Biology. Apterae viviparae outnumber alatae viviparae in March and May 
in Britain but alatae outnumber apterae in June and July. Only apterae viviparae 
are known for September and Cinara abieticola does not seem to have been collected 
in Britain in August. The summer is said to be spent on the roots of Abies in 
special chambers prepared by ants. Oviparae and alate males have been seen from 
Abies pindrow from Pakistan and were described as Lachnus pubescens by Wellenstein 
(1930 : 743-745) from Europe. Heikinheimo (1963 : 185) has recorded natural 
enemies. 

Notes. Cinara abieticola is a large aphid which sporadically occurs in large 
numbers between March and June. Heikinheimo has discussed the possibility 
that Aphis borealis Curtis is the earliest name for abieticola. Borner (1952 : 44) 
has identified shorter haired specimens with Todolachnus confinis (Koch), presumably 
because of the long rostrum mentioned in the original description. Lachnus confinis 
was apparently described from a vagrant alata and so was probably fully pigmented. 
The hind tibiae are figured as being yellow and apart from the length of the rostrum 
the description fits pilicornis. Hottes (1954 : 260) says that C. lasiocarpae is 
very similar to abieticola but that it is smaller and has paler legs, bearing shorter 
and more strongly inclined hairs. Most of the specimens of this group studied have 
been separable by this combination of characters but intermediates in all characters 
occur. Larger specimens from Colorado may have rather dark tibiae bearing 
inclined hairs and the short-haired European forms of abieticola have inclined tibial 
hairs. It seems likely that Cinara vanduzei, C. cecconii, C. pubescens, C. confinis 
(Koch) Borner, C. lasiocarpae and C. abieticola ssp. bulgarica are all forms of abieticola. 
If there really are two or more species, then they are both now probably holarctic 



REVIEW OF CINARA IN BRITAIN 



127 



in distribution. Cinara hattorii Kono & Inouye, 1938 (= konoi Inouye, 1956) 
and C. longipennis (Matsumura, 1917) from Abies in Japan are placed in the subgenus 







12 





Figs 6-12. Cinara acutirostris , aptera vivipara. 6, third antennal segment, x 87. 
7, apex of fifth and sixth antennal segment, x 87. 8, sixth antennal segment, x 450. 
9 & 10, fourth and fifth rostral segments with eight accessory hairs, different views, 
x no. 11, mesosternal tubercle, x no. 12, hind tarsus, x no. 



128 



V. F. EASTOP 



Dinolachnus with abieticola by Inouye (1970 : 88-90) and Cinara sonata Hottes, 
1955, and C. grande Hottes, 1956, from Abies in North America are also said to be 
related to abieticola. 

Aizenberg (1956 : 139) recognized, presumably from the description alone, that 
vanduzei Swain was not piceae Panzer (= grossa). Aizenberg had apparently not 
seen Hottes & Essig (1954 : 95-97), in which vanduzei is placed as a synonym of 
abieticola and concluded that vanduzei was related to Cinara bogdanowi. Pitsaria is 
presumably a lapsus for Pity aria. 

Type-species of Dinolachnus Borner, 1940, as Lachniella cilicica var. cecconii Del 
Guercio. 



Cinara acutirostris Hille Ris Lambers 
(Text-figs 6-13) 

[Cinara montanicola Borner; Stroyan, 1955 : 332-333. Misidentification.] 

Cinara acutirostris Hille Ris Lambers, 1956 : 246-249. [Types colln. D. Hille Ris Lambers, 

described from Netherlands, Czechoslovakia, Italy, England]; Stroyan, 1957 : 355 ; 

Pintera, 1966 : 309. 
Cinaria acutirostris (Hille Ris Lambers) Heinze, 1962 : 166. 

Material examined. 

England: Cambridge, Botanical Gardens, Pinus nigra var. calabrica, 20. iii.1951, 
first instar larvae of fundatrices and egg-breakers; 25.vi.1951, 12 apterae and 2 
alatae viviparae; 2.vii.i95i, 4 apt., 8 al. ; 13.x. 1950, 4 oviparae, 1 apterous male 
(V.F.E.); 9.x. 1950, 18 oviparae, 1 apterous male (H. L. G. Stroyan coll. & colln.), 
except for 6 oviparae in BMNH; Kent, Wye, 40' trap, 3.vii.ig69, 1 al. (N. R. Maslen), 
Forestry Commission Colin. 

France: Beauregard, Pinus nigra, n.v.1966, 2 fundatrices {Laurent coll.), 
D. Hille Ris Lambers colln. Italy: Calabria, Sila, Pinus calabrica, August 1968, 
2 apterae (Tremblay coll.), D.H.R.L. colln. 




Fig. 13. Cinara acutirostris. Fore wing, x 30. 



REVIEW OF CINARA IN BRITAIN I2Q 

Host-plants. Seen from Pinus nigra var. maritima (= calabrica), originally 
described from P. nigra (= austriaca) and also recorded (Ilharco, 19686 : 248) from 
Pinus pinea. 

Distribution. Specimens have been seen from England, France and Italy 
were originally described from the Netherlands and are also recorded from Bulgaria' 
Czechoslovakia and Portugal. 

Biology. Eggs hatch in the second half of March at Cambridge Alatae 
viviparae are produced in the second half of June and the first week in July 
Oviparae and apterous males were collected in October. Cinara acutirostris has 
not been seen from Britain in August or September. When present it may occur 
in large numbers and be the prey of various natural enemies. Recorded in the 
original description from a nest of the sphecoid wasp Psemdus fuscipennis. Stroyan 
( J 955 : 332) records Coccinella quadripuncta as a predator. 

Cinara boerneri Hille Ris Lambers 
(Text-figs 14-18) 

[Aphis laricis Walker, 1848 : 102-103, partim, second variety; 1852 : 957-958 partim 1 
? Lachnus laricis Koch, 1856 : 241-243. [No types. Locality and year not given, presumably 

Germany]: Cholodkovsky, 1898 : 666, partim, nee Walker 1848 
[Lachnus pinicolus (Kaltenbach) Buckton, 1881 : 52-53, partim. Misidentification 1 
? Lachniella laricis cuneomaculata Del Guercio, 1909 : 291-293. [Types unknown. Italy- 

Pratolino, Larix europaea, July, aptera vivipara ] 
[Panimerus laricis (Walker) Theobald, 1929 : i 35 -i39. partim, the alatae viviparae and 

alate males. Misidentification.] 
[Cinara taeniata (Koch) Braun, 1938 : 479, partim. Misidentification ] 
Cinara lancicola Borner, 1939 : 75- [Types, Deutsches Entomologisches Institut. Germany ■ 

Bohmen & Ostmark, Larix]; Stroyan, 1955 : 331 (nee Lachnus laricicolus Matsumura 1917) 
Cinara (Cinarella) laricicola Borner, Borner, 1949 : 59 (Cinarella Borner, 1949, nee Cinarella 

Hille Ris Lambers, 1948). 
Cinara (Cinarellia) laricicola Borner, Borner, 1952 : 41 ; Pasek 1954 ' 250-253 
Cinara boerneri Hille Ris Lambers, 1956 : 246. [Norn. n. pro Cinara laricicola Borner 1939 

nee Matsumura, 19x7]; Stroyan, 1957 : 354~355; Szelegiewicz, 1962 : 80 
Cinara (Cinarellia) boerneri Hille Ris Lambers, Heinze, 1962 : 154, 156; Pintera, 1966 : 

Material examined. 

England: Devon, Exeter, Larix, 14.ix.1965, 1 aptera {H. G. Morgan & colln ) • 
Dorset, Weymouth, Mt Edge, 17.X.1894, 2 alate males (Bignell); three other alate 
males mounted on individual slides in the Buckton collection bear the following 
data, 'Weymouth, October, Bignell', slide 348; 'Weymouth, larch, September^ 
slide 349; 'Weycombe [Surrey], Sept.', slide 350, and probably belong to the same 
series as the previous two males. Buckton (1881 : 53) records rearing winged 
terms presumably at his home, Weycombe, Surrey, from nymphs collected by 
Bignell at Weymouth, Dorset. Hants, Alice Holt Lodge, Larix, 7 .ix.i 9 6 2 , 3 apterae 
(t. I. Carter), Forest Research Station colln. Berks, Tubney, Larix decidua 
28.v1.1956, 3 apt., 1 al. (H. L. G. Stroyan & colln.). Surrey, Kew Gardens Larix 



I 3° 



V. F. EASTOP 



decidua, 25.vii.1964, 1 apt.; 13.viii.1961, 3 apt.; 9JX.1962, 7 apt.; 22. ix. 1962, 
6 apt.; 14.X.1961, 1 ovipara; 26.x. 1968, 1 ovipara {V.F.E.); Fernhurst, Larix 





Figs 14-18. Cinara boereri. 14-16, male genitalia, x 135. 17 & 18, aptera vivipara. 
17, right siphunculus and part of fifth abdominal tergite, x 87. 18, second and third 
antennal segments, x 87. 



REVIEW OF CINARA IN BRITAIN 131 

decidua, 11.vii.1967, 1 apt., 1 alatoid nymph (V.F.E.); Richmond Park, Larix, 
7.vii.i968, 1 apt. (V.F.E.); Wisley, R.H.S. gardens, 11.ix.1966, 1 apt. (K. M. 
Harris & V.F.E.); Kent, Wye, vagrant, June 1928, 1 al. (F. V. Theobald); London, 
Southgate, larch, 11. ix. 1847, x a P r - (F- Walker, 497); 2.ix.i847, 1 larva (495); 
28.vii.1847, 1 al. (F. Walker, 499). Middlesex, Moor Park, Larix decidua, 15.vii.1962, 
1 al. & larvae; 17.viii.1952, 1 apt. (V.F.E.). Herts, Harpenden, Rothamsted 
Experimental Station tower, 9.VI.1968, 1 al. (N. R. Maslen), Forestry Commission 
colln. Cambridge, Botanical Garden, Larix decidua, 10. vi. 1952, 2 apt., 1 alatoid 
nymph (V.F.E.); 12.x. 1950, 2 alate £$ (H. L. G. Stroyan & colln.); Larix dahurica 
var. pendula, 15.viii.1951, 11 apt.; L. sibirica, 15.viii.1951, 1 apt., L. dahurica, 
11. ix. 1952, 3 apt.; L. laricina, 10. x. 1950, 2 apt. (V.F.E.). Suffolk, Bury St Ed- 
munds, Broom's Barn Expt. Stn, 29. vi. 1968, 1 al. (N. R. Maslen), Forestry Com- 
mission colln. Yorks, Malton, High Mowthorpe Experimental Station, 30. vi. 1968, 
1 al. ; 12.viii.1968, 1 al. (A T . R. Maslen), Forestry Commission colln. Lanes, Meathorp 
Wood, larch, 5.ix.i966, 3 young larvae (E. J. White). Westmorland, Windermere, 
larch, 12.viii.1913, 2 ovip. ; 16.viii.1913, 1 <$ (F. V. Theobald). Scotland : Edinburgh, 
Bush nursery, Larix grafts, 22. xi. 1962, 1 al. £ (C. I. Carter, Forestry Comm. colln.). 

Austria: Glochner, Larix europaea, i.viii.1943, 1 apt. (V. Pasek). Czecho- 
slovakia: Banska Stiavnica, L. decidua, 20.viii.1951, 3 apt.; 4JX.1949, 3 apt.; 
Polana, L. decidua, 20.vii.1951, 2 apt. (V. Pa§ek). 'Conifer' without further data, 
1 apt., Theobald colln. Germany: Berlin Dahlem, L. decidua, 8.vii.i95o, 1 apt., 
W. Quednau colln. (K. Heinze). Italy: Zuel, nr. Cortina, Larix, 30.vii.1969, 
5 apt., 1 al. (V.F.E.). Netherlands: Bennekom, L. decidua, 15.x. 1946, 1 apt. 
vivipara, 7 oviparae, 5 alate males (D. Hille Ris Lambers & colln.). 

Host-plants. Specimens have been seen from L. decidua (= europaea), L. 
gmelini (= dahurica), L. laricina and L. sibirica and are also recorded from Larix 
hptolepis. 

Distribution. Specimens have been seen from Austria, Czechoslovakia, England, 
Germany, Italy, Netherlands and Scotland and have also been recorded from 
Bulgaria (Tashev, 1961 : 157), Latvia (Rupais, 1961 : 13-14), Mongolia (Szelegiewicz, 
19636 : 113), Poland, Sweden, Switzerland, Ukraine and with a query from Yugo- 
slavia (Rihar, 1963 : 264). 

Biology. In England apterae viviparae have been collected in June, July 
and September, alatae viviparae from June 30th to August 12th, oviparae and 
alate males in October. Saeman (1966 : 379) gives an account of the biology. 
Alate males have been recorded from June to early August but have only been 
found in October and November in Britain. Attended by Formica rufa (according 
to Weis (1955 : 472) and by other ants (Pintera, 1966 : 319) in the wild but surviving 
in botanical gardens without ant attendance or with only occasional visits from 
Lasius niger. 

Notes. Closely related to Cinara laricicola (Matsumura) from Japan and to 
C. laricifex (Fitch) from America according to Hille Ris Lambers in Inouye (1962 : 
152). Borner (1952 : 42) lists Lachniella laricis cuneomaculata Del Guercio, 1909, as 
a synonym of C. kochiana but the length of the tibiae agree better with C. boerneri. 
Important in apiculture in Central Europe, according to Borner & Franz (1956 : 29). 



132 V. F. EASTOP 

Cinara bogdanowi (Mordwilko) 

? Lachnus pruinosus Hartig, 1841 : 368. [No types. Germany: Berlin], see Stroyan, 1957 ! 

349- 
[Aphis abietis Walker, 1848 : ioo, partim, 1 alata (see Doncaster, 1961 : 13).] 
[Aphis laricis Walker, 1848 : 102, partim, 1st variety (see Doncaster, 1961 : 92).] 
Lachnus bogdanowi Mordwilko, 1895a : 75, 79, 83, 94, 97-98. [Types unknown. Locality 

not given, presumably Poland]; 18956 : 97, 115-118; Cholodkovsky, 1896a : 150; 1898a : 

657- 6 59; 1902 : 6. 
Lachnus piceicola var. viridescens Cholodkovsky, 1896 : 509. [Types unknown. Locality 

not given, presumably U.S.S.R.]; 1898 : 662. 
Neochmosis abietis (Walker) Theobald, 1929 : 362-354, partim. 
Dilachnus radicicolus Wellenstein, 1930 : 739-743, 751. [Types unknown. Locality not 

given, presumably Germany.] 
Cinara bogdanowi (Mordwilko) Borner, 1932 : 568; Braun, 1938 : 479; Szelegiewicz, 1962 : 

79-80; Shaposhnikov, 1964 : 521; Pintera, 1966 : 309-311. 
Cinara radicicola (Wellenstein) Borner, 1932 : 570. 
Cinaropsis (Pityaria) pruinosa (Hartig) Borner, 1949 : 59; 1952 : 43. 
Cinaropsis viridescens (Cholodkovsky) Borner, 1952 : 43. 
Cinaria intermedia Pasek, 1952 : 96, nomen nudum; 1953 : 4-6. [Lectotype, Academy of 

Science, Prague. Czechoslovakia]; 1954 : I 53 _I 55- Syn. n. 
Cinaria borealis (Curtis) Pasek, 1953c : 225-226; 1954 : 216-218. 
? Cinaria (Mecinaria) radicicola (Wellenstein) Aizenberg, 1956 : 139. 
Cinara (Cinaropsis) bogdanowi (Mordwilko) Stroyan, 1957 : 349-351- 
Cinaropsis (Pityaria) bogdanowi (Mordwilko) Heinze, 1962 : 168-169. 
Cinara borealis (Curtis) Pintera, 1966 : 307-308. 
Cinara (Pityaria) bogdanowi (Mordwilko) Canakcioglu, 1966 : 140. 

Material examined. 

England: London, Southgate, Spruce, 3.vii.i847, 1 alata (F. Walker) (slide no. 
7B); larch, 2. i. 1847, 3 apterae (F. Walker) (496); 11.ix.1847, 4 apt. (F. Walker) 
(slide no. 497). Surrey, Wisley, Picea schrenkiana, 6.viii.i953, 1 apt. (/. P. Don- 
caster). Hants, Alice Holt, light trap, 4.vi.i964, 1 al., Forestry Commission colln. 

Belgium: Jupille, Picea excelsa, n.v.1961, 8 alatae (/. Leclerq coll.), H. L. G. 
Stroyan leg. Czechoslovakia: Jakubov, Picea excelsa, 18.V.1950, 2 apterae, 
Gelnica, P. excelsa, 17. vi. 1952, 2 apt., P. excelsa roots, 30.ix.1950, 2 apt., Ruske" 
Peklany, Picea excelsa, 23. vi. 1952, 2 apt., Bianska Stiavnica, P. excelsa, 22.vii.1952, 
2 apt., Vysoka, P. excelsa roots, 16. ix. 1952, 2 apt. (V. Pasek); Kotlina, 18. vi. 1932, 
1 al. (D. Aubertin) (C. borealis (Curtis) of Pasek). Germany; Moor, Fichten, 
27. v. 1934, 2 apt., (Rebeler coll.), K. Heinze leg. Oberhof, P. excelsa, i.x.1937, 

1 ovipara (K. Heinze). Netherlands: Hoenderloo, Picea, 70. vi. 1946, 6 oviparae 
(Elton coll.), D.H.R.L. leg.; Bennekom, P. alba, i.x.1946, 4 oviparae, D.H.R.L. 

2 colln. Poland: Bydgoszoz Jachcice, P. excelsa, 4.viii.i956, 1 larva (H. Szelegie- 
wicz). Without data, ? Poland, 3 apt., 1 alatoid nymph, ex. coll. Mordwilko. 
Turkey: Artrim, Kurukuriin, 1940 m., Picea orientalis, 1 apt.; Istanbul, Maslak, 
100 m, P. abies, 2 larvae, 20. vi. 1964 (H. Szelegiewicz). 

Host-plants. Seen from Picea abies (= excelsa) and more rarely from other 
members of Eupicea such as P. orientalis and P. schrenkiana (= tianshanica) and 
also recorded from P. glauca. 



REVIEW OF CINARA IN BRITAIN 



133 



Distribution. Seen from Belgium, Czechoslovakia, England, Germany, Nether- 
lands, Poland, and Turkey. Also recorded from Austria, Bulgaria, Norway, 
Ukraine, Moldavia, Latvia and Estonia. 

Biology. Occurs on both the branches and the roots of spruce. Alatae vivi- 
parae occur from May until early July. Oviparae occur in October. According 
to Pintera (1966 : 311) the spring generations live on the two year old or older 
twigs in shady positions and alatae are produced in the third generation. The 
progeny of the alatae develop on the roots or at the base of the trunk. Cinara 
bogdanowi is attended by ants which construct special chambers around the roots 
for the aphids. Mating occurs after the sexuales have left the subterranean shelters 
and the oviparae lay eggs on the bark of the younger twigs. Wellenstein (1930 : 
739-743) describes the oviparae and alate males as Lachnus radicicolus. Anholo- 
cyclic overwintering on the roots has been recorded. 

Notes. Said to be important for the production of 'forest honey' in Central 
Europe (Pintera, 1966 : 311). Inouye (1970 : 73-74) regards the Japanese Cinara 
ezoana Inouye, 1936, as a subspecies of C. bogdanowi. 

Cinara brauni Borner 
(Text-fig. 19) 

? Enlachnus nigrofasciatus Del Guercio, 1909 : 316, 324-326. [Types unknown. Italy: 

Firenze, Pinus sylvestris, 1905.] 
Cinara brauni Borner, 1940 : 1. [Types, Deutsches Entomologisches Institut. Central 

Europe, Pinus austriaca]; Szelegiewicz, 1962 : 80; Tashev, 1964 : 172-173; Pintera, 

1966 : 290, 291; Maslen, 1969 : 228. 
Cinara (Subcinara) brauni Borner; Borner, 1949 : 59; 1952 : 41; Pa§ek, 1954 : 175-179. 

Material examined. 

England: Hants, Alice Holt Gardens, suction trap, 20.vii.1968, 1 alata (N. R. 
Maslen), Forestry Commission collection. 




Fig. 19. Cinara brauni, aptera vivipara. Antennal segments I— III, x 150. 



i 3 4 V. F. EASTOP 

Austria: Wien-Schwechat, Pinus nigra, 7.ix.i97o, 4 apt. (W. Quednau), Czecho- 
slovakia: Banska Stiavnika, P. nigra, 7.vii.i952, 3 apterae, 1 al. ; Detra, P. nigra, 
8.viii.i952, 2 apt., 1 al. (V. Pasek). Hungary: Biikk, Fekete-sar, P. nigra, 9.ix.i965, 
2 apt. viviparae, 1 ovipariform apt. (H. Szelegiewicz & colln. 2942). Netherlands: 
Arnhem, Pinus austriaca, 26.viii.1955, 2 apt. (D. Hille Ris Lambers). Poland: 
Bydgoszez-Osied. L., Pinus nigra, 8.viii.i957, 3 apt., 1 al.; Wladystawowo, distr. 
Puck, P. nigra, 8.VU.1960, 3 apt., 3 al. (H. Szelegiewicz & colln.). Turkey: Istanbul, 
Bahcekoy, no m a.m.s.L, P. nigra var. pallasiana, 18.vii.1963, 3 apt., 1 
alatoid nymph; Istanbul, Bayazit, 80 m a.m.s.L, P. nigra, 7.VL1964, 1 apt., 1 al.; 
Bursa-Uludag, Sarialan, 1600 m a.m.s.L, P. nigra var. pallasiana, 22.viii.1965, 
7 apt. (H. Qanakcioglu). 

Host-plants. Pinus nigra (= austriaca), P. nigra var. caramatica (= pallasiana) 
and P. nigra var. maritima (= laricio). 

Distribution. Specimens have been seen from Austria, Czechoslovakia, England.. 
Hungary, Netherlands, Poland and Turkey and are also recorded from Bulgaria, 
Germany and the Crimea. 

Biology. According to Pintera (1966 : 291) it lives on young shoots attended 
by the ant Formica rufa. 

Cinara costata (Zetterstedt) 
(Text-fig. 20) 

Aphis costata Zetterstedt, 1828 : 559. [Types, Entomologische Museum Lund, teste Wahlgren, 

1939 : 3-4, Lappland]; 1840 : 311; Walker, 1848 : 100-102; 1852 : 957. 
? Cinara symphyti Curtis, 1835 : no. 577, page 2. [Types unknown. England: June, alatae] 

(see Walker, 1852 : 957; Hottes, 1949 : 159). 
Lachnus costatus Hartig, 1839 : 645 (new species). [Types unknown. No locality, presumably 

Germany.] 
Schizoneura costata (Hartig) Hartig, 1841 : 367; Walker, 1852 : 1050. 
[Lachnus fasciatus Burmeister; Kaltenbach, 1843 : 160-161; Koch, 1856 : 237-238; Swain, 

1921 : 211-212. Misidentifications.] 
Dryobius cistatus [sic] Walker; Buckton, 1881 : 78. 
Lachnus farinosus Cholodkovsky, 1892 : 74-75. [Types unknown. Without locality, 

presumably Russia, near Leningrad]; 1896a : 145-146; 1898 : 650-654; van der Goot, 

1915 : 395- 

[Lachnus fasciatus (Kaltenbach) Mordwilko, 1895a : 102; 18956 : 104, 139-146.] 

[Lachniella fasciata (Burmeister) Del Guercio, 1909 : 294-296, partim ?, alata only. Mis- 
identification.] 

Lachnus costatus (Zetterstedt) Jackson, 1919 : 164; Theobald, 1929 : 157-161. 

[Panimerus pinihabitans (Mordwilko) Theobald, 1929 : 132-135 partim, the apterae from 
Windermere. Misidentification.] 

Pterochlorus cistatus [sic] (Walker) Buckton; Swain, 1921 : 211. 

Lachniella costata (Zetterstedt) Borner, 1932 : 571; 1952 : 45; Braun, 1938 : 482; Wahlgren, 
*939 : 3 _ 4; Pasek, 1954 : I 95 _I 97: Heinze, 1962 : 176-178; Higuchi & Miyazaki, 1969 : 31. 

Cinara costata (Zetterstedt) Mordwilko, 1933 : 159; 1948 : 201; Inouye, 1937 : 105; Kloft, 
Kunkel & Ehrard, i960 : 166-167; Hottes, 1961 : 41, 43; Szelegiewicz, 1962 : 81; Shaposh- 
nikov, 1964 : 522; Pintera, 1966 : 311— 314. 

Cinara (Lachniella) costata (Zetterstedt) Eastop, 1966 : 528-529. 

Cinara (Cinaropsis) costata (Zetterstedt) Inouye, 1970 : 84-85. 



REVIEW OF CINARA IN BRITAIN 



135 



Material studied. 

England: without further data, 1 alata (F. Walker), slide no. 266. London, 
Southgate, spruce, 5^.1847, 1 aptera, 2 alatoid nymphs (F. Walker), slide 677; 
21.vi.1847, 1 al. (no. 269); 25.vi.1847, 2 al. & 1 immature (270, 271); 30.vi.1847, 
4 apt., 1 al. (272); 3.vii.i847, 1 apt., 2 al. (276); 30.X.1847, 7 oviparae (274, 275, 
277); 'scotch pine', 9. vi.1847, 1 al. (273); 'scotch fir', 18.vi.1847, 1 apt., 2 al. [F. 
Walker), slide no. 273. Kent, Wye, spruce, 5.vii.i9i3, 9 apt.; 19.V.1927, 13 apt.; 
'Finns sylvestris', 5.V.1913, 1 apt., 3 alatoid nymphs (F. V. Theobald), the correct 
data for this sample is probably 'spruce' 27.V.1913, see data for Cinara pinea from 
Wye in May, 1913 on page 157. Herts, Redbourne, Picea abies, 13. v. 1945, 4 apt., 
4 al. ; Picea sp., 13. v. 1945, 8 apt., 4 al. (/. P. Doncaster). Harpenden, Rothamsted 
tower trap, 9.VL1968, 1 al., Forestry Commission colln. Berks, Midgham, Picea 
excelsa, 15. v. 1948, 9 apt. (V.F.E.). Gloucs., Cowley Manor, June 1959, 1 al. (R. S. 
George). Hants, Alice Holt, Picea abies, 14.vi.1967, 3 apt., 1 alatoid nymph (H. C. 
Dale). Somerset, Long Ashton, 1964, 1 al. (A. Stringer). Bucks, Slough, 1937, 
2 apt., 2 al. (H. Downes). Cumberland, Great Salkeld, Pinns sylvestris, 20.vi.1914, 
1 al. (F. V. Theobald), the alatoid nymph on the same slide is Schizolachnus pineti. 
Westmorland, Windermere, spruce, 5.V.1913, 3 immature (F. V. Theobald) ; 13.V.1915, 
1 apt. {R. Roberts), F.V.T. colln. 

Czechoslovakia: Prague, flying, 2.VL1964 (V.F.E.). Greenland: 1929, sto- 
mach of Phalarope, thorax and abdomen of 1 al. (0. W. Richards). Netherlands: 
Bennekom, Picea sp., 21. vi. 1949, 3 apt., 2 al. (D. Hille Ris Lambers & H. L. G. 
Stroyan coll.), H.L.G.S. colln. Japan : Hokkaido, Sapporo, Mt Soranumar, Picea sp., 
3.VU.1964, 3 apt. (H. Takada). Poland: Bydgoszcz- Jacheire, Picea excelsa, 
4.viii.i956, 1 apt. (H. Szelegiewicz). 

Australia: New South Wales, Mittagong, 'Abies excelsa', 25. ix. 1959, 7 apt.; 
Picea coseana glauca', 25.ix.1959, 3 apt. (V.F.E.). Tasmania, Oaklands, Picea, 
22. v. 1952, 2 larvae, Tasmania Dept. Agriculture colln. Canada: Ontario, Ottawa, 
Picea abies, 4.VL1951, 1 apt. (W. R. Richards). 




Fig. 20. Cinara costata. Wings, x 24. 



i 3 6 V. F. EASTOP 

Host-plants. Picea abies (= excelsa) and also recorded (Pintera, 1966 : 312) 
from other Eupicea such as Picea glauca, P. glehni and P. pungens and from P. 
jezoensis in the Cascita group and P. omorika of the Omorika group. 

Distribution. Specimens have been seen from Czechoslovakia, England, 
Greenland (from stomach of Phalarope), Netherlands, Japan, Poland, New South 
Wales, Tasmania and Ontario. Cinara costata has also been recorded from Austria, 
Finland, Germany, Norway, Scotland, Sweden, U.S.S.R., Ukraine, Latvia, Estonia, 
Sachalin, Yugoslavia, California (see 'Notes') and Washington. 

Biology. Numerous apterae and some alatae can be collected in May in England 
and apterae are less common in June. Occasional apterae and alatae have been 
collected in July. Oviparae occur in October. C. costata seems not to have been 
collected in Britain during August or September. According to Pintera (1966 : 
312-313) C. costata lives on the older twigs all the year and is not or only rarely 
attended by ants. In September, 1959 in New South Wales, it was found both on 
the second year twigs and on the lower branches covered with earth, apparently 
by ants. Cholodkovsky (1892 : 74-75) gives an account of the biology, including 
alate males, presumably near Leningrad. 

Notes. 

The type-species of Lachniella Del Guercio, 1909 (as fasciata) 

Lachniella is a real species-group in the sense that there are a number of species 
of similar appearance with similarly pigmented wings, few rhinaria on the antennae 
of the alatae and oviparae with strongly swollen hind tibiae. Otherwise the species 
are similar to Cinaropsis and perhaps Cinaropsis should be regarded as a synonym 
of Lachniella, used as a sub-genus of Cinara. The confusion concerning the use 
of Lachniella resulted from Lachnus fasciatus Burmeister, 1835 being selected as the 
type-species of Lachnus Burmeister by Westwood, 1840, without there being agree- 
ment as to the identity of Lachnus fasciatus. Some authors regarded L. fasciatus as 
a synonym of Lachnus roboris (L.) and others as a synonym of Lachnus costatus. 

Cinara nimbata Hottes, 1954, from North America is similar to costata but bears 
shorter and thicker hairs. It is not clear whether the record of costata from Picea 
sitchensis in California by Hottes & Essig (1953 : 172) really applies to costata or 
to the species subsequently described as Cinara sitchensis Hottes, 1958. 

Cinara cupressi (Buckton) 
(Text-fig. 21) 

Lachnus cupressi Buckton, 1881 146-47. [Types BMNH. England]; Cholodkovsky, 

1898 : 669-670. 
Lachnus juniperinus Mordwilko, 1895a : 75, 79, 82, 94, 102. [Types unknown. Poland]; 

18956 : 103-104, 134-136; Cholodkovsky, 1898 : 668-669. 
Lachniella juniperinum (Mordwilko) Del Guercio, 1909 : 305-306. 
Lachniella tujae Del Guercio, 1909 : 288, 309-310. [Types unknown. Tuja without other 

collection data, presumably Italy.] 
? Lachniella juniperi signata Del Guercio, 1909 : 289, 314-315. [Types unknown. Italy, 

nr Firenze, Thuja occidentalism 



REVIEW OF CINARA IN BRITAIN 



J 37 



191 1 : 541-543; van der Goot, 1915 : 396-399. Mis- 



[Lachnus juniperi (De Geer) ; Essig 

identification.] 
Dilachnns cupressi (Buckton) Swain, 1921 : 212-213. 
Lachnns sabinae Gillette & Palmer, 1924 : 9-1 1. [Types. 

(= Juniperus) scopulorum.] Syn. n. 
Panimerns cupressi (Buckton) Theobald, 1929 : 148-149. 
[Panimerus juniperi (De Geer) Theobald, 1929 : 151 partim 

virginiana at Kew. Misidentiftcation.] 
Panimerus tujae (Del Guercio) Theobald, 1929 : 153-154. 
Cinara sabinae (Gillette & Palmer) Gillette & Palmer, 1931 
Cinara cupressi (Buckton) Borner, 1932 : 570; Braun, 1938 

Szelegiewicz, 1962 : 83. 
Cinara thujae (Del Guercio) Braun, 1938 : 480. 
Neochmosis cupressi (Buckton) Kloet & Hincks, 1945 
Neochmosis tujae (Del Guercio) Kloet & Hincks, 1945 
Cupressobium cupressi (Buckton) Borner, 1952 : 45. 
Cinara canadensis Hottes & Bradley, 1953 : 86-87. 

Ottowa: Ontario, Juniperus virginiana.] Syn. n. 
Cinara (Cupressobium) cupressi (Buckton) Eastop, 1958 : 93. 
Cinara juniper ina (Mordwilko) Ossiannilsson, 1959 : 379. 
Cupressobium juniperinum (Mordwilko) Heinze, 1962 : 174-176 
[? Cinara tujafilina (Del Guercio); Tashev, 1944 : 173. Misidentiftcation.] 



Colorado: Fort Collins, Sabina 



the specimens from Juniperus 



867; Palmer, 1952 : 45. 
480; Hottes & Essig, 1953 



172; 



70. 
70. 

[Type, Canadian National Collection, 



lATERIAL STUDIED. 

England: Cornwall [Probus, cypress], 17. xi. [1879], 4 apterae (/. T. Roscawen) 

type-series of cupressi). Gloucs., Long Ashton, Cupressus sp. ( May- June 1937, 

apt. (C. L. Walton). Somerset, Weston-super-Mare, Cupressus macrocarpus, 




Fig. 21. Cinara cupressi. Male genitalia, x 000. 



138 V. F. EASTOP 

June 1937, 4 apt.; cypress, June 1937, 2 apt. (C. L. Walton); Bristol, Thuja sp., 
7.vii.i9i3, 7 apt., 1 alata (F. V. Theobald). Surrey, Kew Gardens, Juniperus 
virginiana, 23.viii.1919, 4 apt. (F. Laing); Wimbledon, Thuja sp., 5.VL1935, 2 apt. 
(C. T. Gimingham) ; Wisley, Cupressus macrocarpus, 20.ix.1923, 17 apt. (F. V. 
Theobald). Berks, Reading, C. macrocarpus, 16. vi. 1957, 5 apt., 1 al., Eastwood's 
nursery; Cupressus sp., 3.VL1950, 1 apt. (V.F.E.); Ascot, Silwood Park, 40' suction 
trap, 10.viii.1968, 1 al., Forestry Commission Colin. Bucks, Slough, R.C.S. Re- 
search Station, cypress, June 1937, 2 apt., 3 al. (Dowries). Kent, Bedgebury 
Pinetum, Juniperus chinensis, 8.vii.i969, 1 al. ; Wye College, suction trap, 6.viii.i969, 
1 al. (N. R. Maslen). 

Czechoslovakia: Bianska Stiavnica, Biota orientalis, 9. v. 1950, 1 apt. (V. Pasek). 
Germany: Bad Zwischenghr, Juniperus virginiana, 12.V.1959, 2 ? fundatrices 
(Gunkel coll.), K. Heinze leg. Friedhof, Thuja occidentalis, 31.x. 1957, 1 ovipara 
(K. Heinze). Berlin, T. occidentalis, October 1953, 6 ovip., 1 alate male (D.E.J.), 
D. Hille Ris Lambers colln. Netherlands: Nalkenburg Z. 1., T. occidentalis, 
7. v. 1930, 1 apt. (D.H.R.L.); Wageningen, Thuja occidentalis ohlendorffi, 19. vi. 1954, 
29 apt., 5 al. (D.H.R.L. & J. P. Doncaster). T. occidentalis, 2 ovip., 2 al. males, 
October, 1940 (D.H.R.L.). Poland: Olsztyn, T. occidentalis, 18 apt. (S. Huculak). 
Turkey: Bacekoy, Istanbul, Cupressus goveniana, 10.iii.1966, 7 apt. (H. Canak- 
cioglu) . 

Canada: British Columbia, Kelowna, Juniperus scopulorum, 5.VL1962, 1 apt. 
(G. A. Bradley). U.S.A. : Penn., Waddle, The Rock, Juniperus virginiana, 13.vi.1965, 
4 apt., 1 al. ; 7.ix.i964, 3 apt. (/. 0. Pepper). 

Host-plants. Thuja occidentalis, Cupressus goveniana, C. macrocarpa, Juniperus 
virginiana and occasionally Juniperus scopulorum and Thuja orientalis. 

Distribution. Specimens have been seen from Czechoslovakia, England, 
Germany, Netherlands, Poland, Turkey, British Columbia and Pennsylvania, and 
are also recorded from Iraq, Italy, Latvia, Sweden, Ukraine and Ontario (original 
description of canadensis). 

Biology. Apterae viviparae live on the aerial parts of the hosts from May to 
September and were described from November in Cornwall where the winter is 
mild. Alatae viviparae are produced between mid June and mid August and 
oviparae and alate males in October. Gunkel (1963a : 1-48 & 1963& : 329-341) 
has given an account of the natural enemies and population dynamics of Cinara 
cupressi. 

Notes. Takahashi's record (1941 : 150) of C. cupressi from Cupressus torulosa in 
Victoria was most probably based on specimens of Cinara fresai. Lachnus juniperi- 
nus seems to be a synonym of Cinara cupressi but Eastop (1966 : 528) and perhaps 
Braun (1938 : 480) applied the name Cinara juniperina (Mordwilko) to C. fresai. 

The apparently discontinuous host-plant preferences have led to a number of 
synonyms. Specimens from Juniperus virginiana have been described as Cinara 
canadensis and from /. scopulorum as Cinara sabinae. 



REVIEW OF CINARA IN BRITAIN 139 

Much of the description of Cinara difficilis Hottes & Frison, 1931, from Juniperus 
virginiana, agrees with C. cupressi but the hairs on the body and appendages of 
difficilis are shorter than in cupressi and the ultimate rostral segment of difficilis 
bears about 20 accessory hairs distributed over both surfaces. 



Cinara escherichi (Borner) 

[Lachnus nudus (De Geer) Mordwilko, 1895a : 78, 79, 81, 82, 99; 18956 : 100, 119-124; 

Cholodkovsky, 1898 : 635, 642-643. Misidentiftcations.] 
? Lachniella picta Del Guercio, 1909 : 293-294. [Types unknown. Italy: Monte Boni, 

Pinus sylvestris, 27. v. 1905, aptera.] 
Cinara nuda (Mordwilko) Hottes, 1930 : 186-187; Pintera, 1966 : 314. 
Cinaria escherichi Borner, 1950 : 2. [Types Deutsches Entomologisches Institut. Austria: 

Burgenland]; 1952 : 43; Pasek, 1954 : 148, 149-151; Heinze, 1962 : 162. 
Cinara escherichi (Borner) Kloft, Kunkel & Ehrhardt, i960 : 165; Szelegiewicz, 1962a : 81. 

Material studied. 

England: Hants, Alice Holt, suction trap, 24. vi. 1967, 1 alata (C. /. Carter & 
N. R. Maslen), Forestry Commission colln. 

Czechoslovakia: Jakubov, Pinus sylvestris, 19. i. 1950, 2 fundatrices; Banska 
Stiavnica, P. sylvestris, io.v.1950, 1 aptera; 7^.1952, 2 al. ; 3.viii.i952, 2 apt. 
(V. Pasek). Poland: Bydgoszcz- Jachcice, Pinus sylvestris, 12.viii.1957, 3 apt., 
3 al. ; Stamirowice distr., Grojec, P. sylvestris, 18.viii.1961, 4 apt. (H. Szelegiewicz 
& colln.). 

Host-plant. Pinus sylvestris. 

Distribution. Originally described from Austria, specimens have been seen 
from Czechoslovakia, England and Poland, and have also been recorded from 
Germany. 

Biology. According to Pintera (1966 : 314) Cinara escherichi lives in large 
colonies on the trunk or basal parts of the older branches in the spring and in rather 
smaller colonies or individually under the bark during the summer. The colonies 
are attended by ants. 

Notes. Cinara escherichi is a member of the 'pitii-gvoup', generally similar to 
C. acutirostris but bears more numerous but shorter hairs. Authors after Mordwilko, 
1895, and before Szelegiewicz, 1962, who refer to Cinara nuda, were applying the 
name to C. pini (L.) or to one of its subspecies. Cinara canatra Hottes & Bradley 
from Pinus banksiana in North America is similar to C. escherichi except for the 
pigmented dorsum of C. canatra and that the second antennal segment of the summer 
apterae viviparae of C. escherichi bear 9-13 hairs, while the second antennal segments 
of C. canatra bear 5-10 hairs. The eighth abdominal tergite of C. canatra bears 
10-14 hairs while the eighth tergite of the summer apterae of C. escherichi bears 
14-23 hairs. 



140 V. F. EASTOP 

Cinara fresai Blanchard 

[? Cinara juniperina (Mordwilko) Braun, 1938 : 480. Misidentification.] 

Cinara fresai Blanchard, 1939 : 860-862. [Types Institute de Sanidad Vegetal, Buenos 
Aires; Argentina: Tandil, Cupressus macrocarpa, alata.] 

[Cinara cupressi (Buckton) ; Takahashi, 1941 : 150. Misidentification.] 

Cinara wacasassae (Tissot, 1945 : 49-52. [Types Florida Agricultural Experiment Station 
Gainsville; U.S.A.: Florida.] Syn. n. 

[Cinara juniperi (De Geer) ; Cottier, 1953 : 77-81. Misidentification.] 
? Cupressobium mordwilkoi Pasek, 1954 : 3°6-3°9; Heinze, 1962 : 174. 

[Cinara {Cupressobium) cupressi ? juniperina (Mordwilko) Eastop, 1961 : 75. Misidentifica- 
tion.] 
? Cinara mordwilkoi (Pasek) Szelegiewicz, 1962 : 84. 
? Cinara {Cupressobium) mordwilkoi (Pasek) Shaposhnikov, 1964 : 524. 

Cinara maui Bradley, 1965 : 668-670. [Types Forest Entomology Laboratory, Winnipeg; 
Hawaii : Cryptomeria japonica, apterae.] Syn. n. 

[Cinara {Cupressobium) juniperina (Mordwilko) Eastop, 1966 : 528; Mound, 1969 : 62. 
Misidentification.] 

Material studied. 

England: Surrey, Farnham, Alice Holt Research Station, Juniperus sabina 
var. tamariskifolia, 25. vi. 1956, 2 apterae and alatoid nymph {D. Bevan coll.), via 
C. I. Carter. Ham, Juniperus horizontalis, 2-31. x. 1966, 27 apt., 3 alatae (L. A. 

Mound) . 

Australia: New South Wales, Wentworth Falls, Cupressus arizonica, 30. i. 1958, 
8 apt. (K. M. Moore) (4 in N.S.W. Forestry Dept. colln.) ; Elizabeth Bay, Cupressus 
sp., 21.1.1963, 12 apt. {Webster coll.), via K. M. Moore; Collaroy, Cupressus brunniana, 
8.^.1962 (G. P. Wright coll.), via K. M. Moore; Mittagong, Cupressus torulosa, 
25. ix. 1959, 2 alatae and one first instar larva (V.F.E.). A.C.T., Canberra, flying, 
22.viii-12.ix. 1959, 3 al. ;? Cupressus sp., 1-5. ix. 1959, 4 apt., 3 al., Cupressus sabina, 
3-10. ix. 1959, 5 apt., 7 al.; 3.x. 1959, 2 apt. & alatoid nymphs. Juniperus sp., 
6.1X.1959, 6 al., 6 larvae. 'Juniper or cypress', 3.ix.i959, 3 apt. Cupressus sp., 
16. xi. 1959, 8 apt. {D. Goodchild) (4 apt. in C.S.I.R.O. colln.). Tasmania, Claremond, 
cypress, 12.viii.1957, 2 apt., 3 al. ; Margate, Cupressus macrocarpa, 23.xii.1964, 
5 apt. (E. J. Martyn). Victoria, Melbourne, Brighton Beach, Cupressus sp., 
2.V.1959, 13 apt., 2 al. (4 apt. now in colln. A. Pintera) ; 21-25.vi.1959, 6 apt., 
Melbourne, Cupressus lambertiana, near ends of twigs, 26. v. 1959, 3 apt. ? Cupressus, 
8.vii.i959, 4 apt. (V.F.E.). Maffra, Mt. Lithgow, 14.iv.1938, 2 apt., Victoria Dept. 
Agric. 

Chile: Plazoleta Yungue, Mas A Tierra, Islas Juan Fernadez, Cupressus sp., 
7.iii.i968, 2 apt. (C. W. O'Brien coll., leg. & 1 in colln A. G. Robinson). 

U.S.A.: California, Berkeley, Juniperus ? scopulorum, 23.iii.1964, 1 apt., 3 al. 
(V.F.E.). San Diego, Juniperus sp., 30.iii.1964, 2 al. & larvae (R. C. Dickson, 
0. Heie & V.F.E.). Florida, St Augustine, Juniperus silicicola, 19.iv.1945, 1 apt., 
1 al. {A. N. Tissot) (metatypes of C. wacasassae). Oregon, Portland, meyer juniper, 
8. i. 1963, 2 apt. (F. P. Larson coll.), G. F. Knowlton leg. 



REVIEW OF CINARA IN BRITAIN 141 

Host-plants. Juniperus horizontalis, J. sabina, J. silicicola and more rarely 
/. chinensis, J. scopulorum and /. squamata; Cupressus macrocarpa, C. torulosa 
and more rarely C. arizonica, and recorded from Cryptomeria japonica as Cinara 
maui. 

Distribution. Specimens have been seen from England, New South Wales, 
Tasmania, Victoria, Chile, California, Florida and Oregon and have also been 
recorded from New Zealand. 

Biology. Alatae viviparae have been found in both June and October in England. 
Sexuales are unknown, unless Cinara mordwilkoi is a synonym. An aphid of sporadic 
appearance, it was common in Melbourne, Victoria from May to July 1959 and at 
Canberra, A.C.T. from August to December 1959, but has apparently not been 
common there since (R. D. Hughes, personal communication). Cinara fresai lives 
under the older branches with ants and alatae viviparae are produced in large 
colonies. 

Notes. Braun (1938 : 480) keys out Cinara juniperina as having a body-length 
of more than 3 mm, the fifth antennal segment as long as the sixth and living on 
Juniperus. He does not give locality data. Pasek's (1954 : 306-309) description 
of Cinara mordwilkoi may also apply to Cinara fresai but it is not likely since Pintera 
and Szelegiewicz found mostly oviparae while fresai is known only from viviparae. 
If C. mordwilkoi is distinct from fresai, then Braun's (1938) record of juniperina 
probably applies to mordwilkoi. Schouteden (1906 : 203) records Lachnus ? 
juniperinus from Juniperus communis in Belgium. The first European specimens 
seen during this study were collected in 1956. 

Cinara fresai is almost unique in Cinara in that the radial sector of the fore wing 
often does not reach the wing apex ; this character is well illustrated in the original 
description. In occasional specimens of C. fresai the radial sector does reach the 
wing apex in the typical Cinara manner and in occasional specimens of C. tujafilina 
the radial sector does not quite reach the wing apex . 

Cinara juniperi (De Geer) 

Aphis juniperi De Geer, 1773 : 56-58. [Types unknown. Locality not given]; 1780 : 38; 

1783 : 77; Fabricius, 1781 : 388; 1794 : 218. 
Lachnus juniperi (De Geer) Kaltenbach, 1843 : 153-154; Koch, 1856 : 243-244; Mordwilko, 

1895a : 101; 18956 : 103, 136-139; Cholodkovsky, 1898 : 667-668; 1902 : 7. 
Lachniella juniperi (F.) Del Guercio, 1900 : 108; 1909 : 312-314; Jackson, 1919 : 164. 
Dilachnus juniperi (De Geer) Swain, 1921 : 213. 
Cinara juniperi (De Geer) Borner, 1932 : 57; Mordwilko, 1933 : 169; Mimeur, 19366 : 253; 

Braun, 1938 : 480; Hottes, 1955 : 103; Szelegiewicz, 1962 : 82. 
Neochmosis juniperi (De Geer) Kloet & Hinks, 1945 : 70. 
Cupressobium juniperi (De Geer) Borner, 1952 : 44; Pasek, 1954 : 304, 305-306; Heinze, 

1962 : 174. 
Cinara {Cupressobium) juniperi (De Geer) Eastop, 1958 : 93; Shaposhnikov, 1964 : 524. 
[? Cinara mordwilkoi (Pasek) Szelegiewicz, 1962 : 84. Misidentification.] 

Material studied. 
England: London, Southgate, juniper, 4^.1847, 7 apterae, 4 alatae (F. Walker) 



I 



142 V. F. EASTOP 

(slide nos. 459 & 560); 25. vi. 1847, 7 apt., 1 al. (461); 21.viii.1847, 2 apt. (463); 
30.x. 1847, 3 apt. (F. Walker), 462. Kent, Wye, juniper, May 1913, 3 al., June 
1913, 1 apt. (F. V. Theobald). From Juniperus communis, Trottiscliffe Park, 
August, 1968, 1 apt. ; Crookhorn Wood, November 1968, 2 apt. ; Shoreham, White 
Hill, January 1969, 1 apt. (L. K. Ward). Surrey, Wisley, Juniperus communis, 
30.vii.1953, 11 apt. (/. P. Doncaster) ; /. communis effusa, 22. v. 1968, alatoid nymphs 
(K. M. Harris); J. communis hibernica, 14.V.1968, 11 apt., 2 al. & many alatoid 
nymphs; /. communis nana, 22.V.1968, 1 al. & alatoid nymph; /. chinensis kewensis, 
22.V.1968, 5 alatoid nymphs, 4.11.1969, 9 apt. (K. M. Harris). Kew Gardens, 
J. chinensis, 26. ix. i960, 7 apt. (V.F.E.). Wimbledon, 'sallow catkins', 16. v. 1930, 
1 al. (C. N. Hawkins); Mickleham Down, juniper, 2.VL1925, 3 apt. (F. Laing). 
Mickleham Common, juniper, 26. xi. 1922 (W. E. China). From Juniperus communis, 
Riddlesdown, 7.VI.1968, 2 al.; January 1969, 2 apt. Newlands Corner, July 1968, 

1 apt. Walton Down, August 1968, 2 apt. ; January 1969, 1 apt. ; Park Down, 
October 1968, 1 apt.; Hackhurst Down, January 1969, 5 apt. (L. K. Ward). Berks, 
Streatly, Green Hill, /. communis, July 1969, 2 apt. (parasitized) ; October 1969, 

2 apt. (parasitized) ; Hogtrough Bottom, /. communis, April 1969, 2 apt. ; May 
1969, 3 al.; July 1969, 1 apt. (parasitized) (L. K. Ward). Oxon, Ewelme Park, 
May 1969, 1 al. ; Ewelme Down, July 1969, 2 apt. (parasitized) ; September 1969, 
2 apt.; Aston, Rowant, July 1969, 2 apt.; Nuffield Common, June 1969, 1 apt., 

1 al.; July 1969, 1 apt.; Peppard Common, April 1969, 1 apt.; July 1969, 2 apt. 
(parasitized); October 1969, 2 apt. ; Bix Bottom, July 1969, 1 apt. (L. K. Ward). 
Bucks,/, communis, Bledlow, 25.ix.1919, 2 apt. (F. Laing). Whiteleaf, 5.viii.i95i, 

2 apt. (V.F.E.). High Wycombe Hill, May 1969, 1 al. ; West Wycombe Hill, April 
1969, 5 apt., 1 brachyptera; May 1969, 3 apt., 3 al. (L. K. Ward). Herts, Harpenden, 
Rothamsted trap F.i, 22. v. 1943, 1 al. (/. P. Doncaster). Rothamsted tower trap, 
31.V.1968, 1 al., Forestry Commission colln. From Juniperus communis, Aldbury, 
11.viii.1944, 4 apt. (J.P.D.); Aldbury Hill, July 1969, 4 apt. (L. Ward); Gustard 
Wood nr. Harpenden, 30. iv. 1957, 4 apt. (H. L. G. Stroyan & colln.); Albury Hill, 
October 1969, 1 apt.; Commonwood Common, May 1969, 2 apt., 4 al. (L. K. Ward). 
Cambridge, Juniperis communis, University botanical garden, 9. v. 1952, 16 apt., 
21.vi.1951, 1 apt. (V.F.E.); 9JV.1954, 1 apt. {H.L.G.S. & colln.). Fleam Dyke, 
May 1969, 3 apt., 2 al., 2 alatoid nymphs; July 1969, 3 apt.; October 1969, 3 apt. 
(L. K. Ward). Northumberland, Alnwick & Wooler, /. communis, May (prior to 
1881), 2 al. (Hardy), G. B. Buckton slide no. 232. Berwick, 2 apt. (prior to 1881), 
G. B. Buckton, 233. Cheviott, May (prior to 1881), 2 al. (Hardy), G. B. Buckton, 
234. Scotland: Sutherland, Brora,/, communis, 11. v. 1936, 2 apt. (G. D. Morrison). 
Perthshire, Trossachs, juniper, 28. vi. 1932, 1 apt., 3 al. (W. H. T. Tarns). Pitlochry, 
juniper, September 1920, 7 apt. (F. V. Theobald). Kincardine, Banchorry, Juniperus, 
5.VL1964, 8 apt., 1 alatoid nymph (L. A. Mound). Inverness, Juniper, July 1933, 
1 apt. (G. D. Morrison). Nethybridge, /. communis, 6.viii.i940, 1 apt., 1 al. (G. D. 
Morrison). Banffshire, Mortlach, 14.vii.1938, 2 apt. (G. D. Morrison). Aberdeen, 
Glen Gairn, 17.viii.1940, 3 immature (G. D. Morrison). Wales: Glams., Gower 
Slade Cliffs, limestone, /. communis, 18. v. 1964, 1 apt., 7 al. (H. L. G. Stroyan & 
colln.). 



REVIEW OF CINARA IN BRITAIN 143 

Azores: Pico, N. slopes, Juniperus oxycanus, 9-17. vii. 1929, 5 apt., 4 al. (/. 
Balfour-Browne). Guernsey: Couture, Juniperus, 11.vi.1951, 15 apt., 2 alatoid 
nymphs (V.F.E.). Czechoslovakia: Banska Stianvika, /. communis, 5. v. 1950, 
3 fundatrices (V. Pasek). Germany: Naumberg, /. communis, summer 1932, 
1 apt., 1 alatoid nymph; Wittental, /. communis, 15. vi. 1961, 3 apt. (K. Heinze). 
Wildberg, (Black Forest), J. communis, 27. vii. 1969, 4 apt. (V.F.E.). Majorca: 
Clan Picafort, 4. v. 1969, 2 apt. (D.J. Williams). Netherlands: Wageningen-Hoog, 
J. communis, 1. v. 1938, 3 apt., ? fundatrices (D. Hille Ris Lambers). Norway: 
As, Vollebekk, trapped, 7. vii. 1954, 1 al. (H. Tambs-Lynche). Switzerland: 
Zenegger Wallis, J. communis, 21.V.1947, 2 apt., ? fundatrices (Stager coll.), D. Hille 
Ris Lambers colln. Turkey: Bolu-Aladag ormani, 1600 m, Juniperus nana, 
3. vii. 1964, 33 apt., 1 al. (H. Canakcioglu). 

New Zealand: Auckland, Juniperus, 11-12.1.1960, 59 apt., 1 alatoid nymph 
(V.F.E.). 

Canada: Seattle, juniper, 29.iii.1940, 1 apt. (C. L. Richie). 

Host-plants. Juniperus chinensis including kewensis, J. communis including 
depressa, effusa, hibernica and nana, rarely /. oxycanus and has also been recorded 
from /. oxycedrus, J. rigida and /. squamata. 

Distribution. Cinara juniperi has been seen from the Azores, Czechoslovakia, 
England, Germany, Guernsey, Majorca, Netherlands, Norway, Scotland, Switzerland, 
Turkey, Wales, New Zealand and 'Canada, Seattle'. Also recorded from Austria, 
Belgium (Schouteden, 1906 : 203), Bulgaria (Szelegiewicz, 1962a : 48), Greenland 
(Hille Ris Lambers, 1952 : 127), Latvia (Zirnits, 1927 : 251), Morocco, Poland, 
Ukraine, Bokhara (Jakhontov, 1929 : 15), Sweden, Japan (on Juniperus rigida: 
Inouye, 1970 : 89-91), Taiwan (on Juniperus squamata: Takahashi, 1937 : 2-3), 
Minnesota (Oestlund, 1922 : 118) and Ontario (MacNay, 1953 : 4). 

Biology. Cinara juniperi is anholocyclic in Southern England and Wales 
with alatae viviparae found only in May and June. Apterae viviparae are common 
in May and June and occur more sparsely on the aerial parts of Juniper during the 
rest of the year. Alatae are also found in Scotland in June but a single alata col- 
lected in August has also been seen. It seems that Cinara juniperi is holocyclic 
in Central Europe. C. juniperi may be a widespread anholocyclic form of a more 
restricted species, holocyclic on Juniperus communis, for which the name C. mord- 
wilkoi (Pasek) may be available. Rupais (1961 : 17-18) records Cupressobium 
mordwilkoi from Latvia. 

Cinara kochiana (Borner) 
(Text-figs 22-25) 

[Aphis laricis Walker, 1848 : 102-103, partim, oviparae and probably '1st var.' (see Doncaster, 

1961 : 93); 1852 : 957.] 
? Lachnns laricis Koch, 1856 : 241-243 partim. [No types, locality not stated, presumably 

Germany]; Cholodkovsky, 1898 : 666 partim. 
' Lachniella laricis cuneomaculata Del Guercio, 1909 : 291-293. [Types unknown. Italy: 

Pratolina, Larix europaea, July, aptera.] 



i 4 4 V. F. EASTOP 

Cinara laricis (Koch) Braun, 1938 : 482, 483, 488-491, ? partim. 

Cinaria kochiana Borner, 1939 : 76. [Said to be nom. n. pro laricis Koch nee Walker, 1848, 

but probably not Koch's species. Borner 's material in Deutsches Entomologische Institut. 

Neither Borner nor Koch gave locality data but both probably from Germany or Austria.] 
? Cinara kochi Inouye, 1939 : 138-141. [Said to be a nom. n. pro laricis Koch, 1857 nee Walker, 

1848, as interpreted by Braun (1957 : 488-491), but Inouye describes five oviparae from 

Japan: Hokkaido, Larix kaempferi, 13. xi. 1936. Types Govt. Forest Experiment Stn., 

Hokkaido.] 
Cinaria (Laricaria) kochiana Borner; Borner, 1949 : 59. 
Laricaria kochiana (Borner) Borner, 1952 : 42; Pasek, 1954 : 2 47~ 2 56; Heinze, 1962 : 160, 

162. 
Cinara (Laricaria) kochiana (Borner) Stroyan, 1957 : 348-349. 
Cinara kochiana (Borner) Hille Ris Lambers, 1956 : 246; Kloft, Kunkel & Ehrhardt, i960 : 

161-164; Szelegiewicz, 1962 : 83; Shaposhnikov, 1964 : 523; Pintera, 1966 : 316-318. 

Material studied. 

England: London, Southgate, larch, 16. x. 1847, 3 oviparae (F. Walker), slides 
500 & 501. 

Czechoslovakia : Banska Stiavnika, Larix decidua, 7^.1952, 2 apterae viviparae 
(V. Pasek), D. Hille Ris Lambers colli).; 23.vi.1952, 1 apt. (V. Pasek), BMNH. 
Netherlands: Putten, Larix leptolepis, 11.viii.1952, 2 apt. (Elton coll.), D.H.R.L. 
colln. Bennekom, L. decidua, 19.x. 1946, 1 ovip. ; Arnhem, L. leptolepis, 29.x. 1952, 
2 ovip.; November 1952, 1 ovip., 1 alate male (D. Hille Ris Lambers & colln.). 
Poland: Chetmowa Gora pow. Kieke, Larix polonica, 20.X.1961, 7 apt., 1 al. (H. 
Szelegiewicz & colln.). Sweden: Brunnhy, Kullen, Larix decidua, 25.viii.1964, 
1 apt. (F. Ossiannilsson & colln.). Korea: Mt Chiri, Pinus koraiensis, 15. vi. 1963, 
1 apt. (Woon Hah Paik). 

Host-plants. Larix decidua (= europaea, including polonica) and L. leptolepis 
(= kaempferi Sarg.) and also recorded from Pseudolarix amabilis (= kaempferi 
Gord.). 

Distribution. Specimens have been seen from Czechoslovakia, England, 
Netherlands, Poland, Sweden and Korea (subsp. kochi see Paik, 1965 : 18-19) 
and are also recorded from Austria, Germany, Italy, Rumania, Ukraine and the 
sub-species kochi from Japan. 

Biology. According to Pintera (1966 : 318) Cinara kochiana lives at the bases 
of the older branches near the trunk, on the trunk or on thick roots. It forms 
large colonies in crevices in the bark and is always associated with ants. Oviparae 
are produced in October and November and an alate male has been seen from 
Arnhem in November. The few alatae seen or recorded were collected in August 
and October. 

Notes. The hairy ultimate rostral segment separates kochiana from other 
European Cinara but several North American species living on Pinus, including 
Cinara pinata, C. piniradicis, C. puerca, C. tanneri and C. thatcheri bear 20-50 
accessory hairs on the ultimate rostral segment. 

The oviparae of C. kochiana bear numerous rather indistinct pseudosensoria 
on the hind tibiae and the eggs in the abdomen are about 2 mm long and 0-9 mm 
wide. 



REVIEW OF CINARA IN BRITAIN 



145 



Braun (1938 : 482-491) found Cinara kochiana and called it C. laricis (Koch), 
although part at least of Koch's description does not fit kochiana. Subsequent 
authors proposing new names for laricis Koch nee Walker or Hartig, have been 




22 



23 





25 




Figs 22-25. Cinara kochiana, aptera vivipara. 22, third antennal segment, x 87. 
23, sixth antennal segment, x 87. 24, processus terminalis, x 450. 25, fourth and 
fifth rostral segments, x 87. 



i 4 6 V. F. EASTOP 

accepting Braun's (1938) interpretation of lands Koch. Cinara kochi Inouye, 
1939 does not seem to be more than subspecifically distinct from kochiana according 
to Hille Ris Lambers (1956 : 246) and Inouye (1970 : 68-69). Borner (1939) 
lists laricis cuneomaculata as a synonym of kochiana but some of the proportions 
given by Del Guercio (hind tibia/hind tarsus 2 = 6-3; rostral 4 + 5/hind tarsus 
2 = 074; rostral 4/hind tarsus 1 = 1-2), agree with boerneri and would exclude 
kochiana. 



Cinara laricis (Walker) 

Aphis laricis Walker, 1848 : 102-103, partim. [Lectotype BMNH, England.] 
? Aphis tenuior Walker, 1849 : xlix. [Possible type in BMNH, described without locality or 

date: see Doncaster, 1961 : 132.] 
? Lachnus laricis Koch, 1856 : 241-243, partim. [No types, locality not given, presumably 

Germany.] 
[Lachnus pinicolus Kaltenbach; Buckton, 1881 : 52—53, partim, the alate males recorded as 

'winged viviparous females'.] 
Lachnus laricis (Walker) Cholodkovsky, 1898 : 666-667; 1902 : 7; Schouteden, 1906 : 203- 

204; van der Goot, 191 5 : 399-400. 
Lachnus maculosus Cholodkovsky, 1899 : 469-470. [Types unknown. Described without 

locality, presumably near Leningrad.] 
Lachniella nigrotuberculata Del Guercio, 1909 : 288, 306-309. [Types unknown. Italy: 

Protolina, Larix sp., spring 1907, apterae and alatae viviparae.] 
[? Lachnus laricifex (Fitch) Patch, 1912 : 164. Misidentification.] 
Lachniella laricis (Walker) Jackson, 1919 : 164. 

Dilachnus laricis (Walker) Swain, 1921 : 213; Wellenstein, 1930 : 749-750. 
Lachnus muravensis Arnhart, 1927 : 471. [Possible types in BMNH; Austria.] 
Panimerus laricis (Walker) Theobald, 1929 : 135-139, partim, the sexuales and some of the 

alatae viviparae are C. boerneri. 
? Cinara laricis (Hartig) Hottes & Frison, 1931 : 155-156. 
Cinara laricis (Walker) Hille Ris Lambers, 1931 : 3; Borner, 1932 : 569; Mordwilko, 1933 : 

169; Braun, 1938 : 478; Pasek, 1954 : 2 53 _2 56; Inouye, 1956 : 216; Doncaster, 1961 : 92- 

94; Szelegiewicz, 1962 : 83; Shaposhnikov, 1964 : 523. 
Neochmosis laricis (Walker) Hille Ris Lambers, 1935 : 63. 

Cinara laricis (Hartig) Inouye, 1962 : 147-151; 1970 : 65-66; Pintera, 1962 : 293-295. 
Cinaria laricis (Walker) Borner, 1939 : 76; 1952 : 42; Heinze, 1962 : 163, 164. 
Cinara doncasteri Pasek, 1953c : 222, 223; 1954 : x 34- I 4 I < J 4 2 - [Holotype, BMNH. Scot- 
land]; Heinze, 1962 : 159, 160; Pintera, 1962 : 292-293. Syn. n. 

Material studied. 

Lectotype aptera vivipara, England: London, Southgate, larch, 2.viii.i847, 
F. Walker slide no. 498. 

One larva on the type-slide and another, 2.ix.i847, F. Walker 494. Surrey, 
Kew Gardens, Larix decidua, 26.x. 1968, 2 oviparae, 1 alate male (V.F.E.). Berks, 
Mortimer, 'Larch/grass area', 2.vii.i959, 1 al., Forest Research Stn, Alice Holt 
colln. Herefordshire, Mortimer, Larix eurolepis, 14.il. 1959, 1 al. (C. /. Carter). 
Derbys, Smerrill Grange, L. europaeus, 22. vi. 1946, 12 apt., 4 al. ; Wensley, L. 
europaeus, 30. vi. 1946, 2 al. (/. P. Doncaster). Westmorland, Windermere, Bishop's 
Wood, Larix leptolassa, 4.VL1914, 3 al. (Rymer Roberts), F. V. Theobald colln.; 



REVIEW OF CINARA IN BRITAIN 147 

Farlwood, L. leptolepis, 13.V.1915, 1 al.; Ellerbeck, Kendal, oak, 26.vi.1914, 1 al. 
(F. V. Theobald). Cumberland, Dunnerdale, L. enropaeus, 5.vii.i953, 5 apt. (/. P. 
Doncaster). Scotland: '48-121' without further collection data, 1 al. (F. Doubleday) , 
Walker colln. no. 676. Inverness, summit of Braeriach, 4000', 28.vi.1932, 2 al. 
(B. M. Hobby) (types of Cinara doncasteri). Perth, Pitlochry, Faskally, Larix 
decidua, 21. v. 1966, 2 apt. (C. /. Carter). Edinburgh, L. decidua, 22.x. 1962, 1 ovi- 
para; 'Bush nursery, Larix grafts', 22.x. 1962, 4 alate males (C. /. Carter) (2 in Forest 
Res. Stn, Alice Holt colln.). 

Austria: 'August 1927, Lachnus muravensis, Dr Arnhart', 5 apterae & larvae 
without further data. Czechoslovakia: 'conifer', 1 al. without further data 
ex F. V. Theobald colln. Banska Stiavnika, Larix decidua, 13. vi. 1950, 2 apt.; 
Bohemia, C. Krumlov, 13.viii.1951, 4 apt. (V. Pasek). Germany: Roth b. Berchtes- 
gaden, L. europea, 23.vii.1952, 6 larvae (H. Schmntterer coll.), K. Heinze leg. Berlin, 
Dahlem, L. europaea, 1 apt., 1 larva, 8.vii.i950 {W. Quednau coll.), K. Heinze leg. 
Italy: Zuel near Cortina, Larix sp., 30.vii.1969, 1 apt., 24 al. (V.F.E.). Nether- 
lands: Arnhem, L. leptolepis, November 1952, 2 oviparae (J.T. B.O.N. ), D. Hille 
Ris Lambers colln. 

Host-plants. Larix decidua (= europaea including polonica), L. leptolepis 
(= kaempferi Sarg.) and also recorded from Larix laricina, L. sibirica and Pseudolarix 
amabilis (— kaempferi Gord.). 

Distribution. Specimens have been seen from Austria, Czechoslovakia, England, 
Germany, Italy, Netherlands, Scotland and have also been recorded from Latvia 
(Rupais, 1961 : 14), Poland, Sweden, Switzerland (Werder, 1934 : 15), Ukraine, 
Mongolia (Szelegiewicz, 1963 : 113), Japan (Inouye, 1962 : 150) and with a query 
from Eastern Siberia (Grechkin, 1962 : 706). Records from America probably 
apply to other species. 

Biology. Apterae viviparae can be collected from May until September and 
alatae viviparae in June and July. Single alatae have also been seen from February 
and May in Western Britain. Oviparae are produced in October and November. 
Cinara laricis lives under the old branches of Larix together with Formica rufa and 
when disturbed the alatae often do not fly but run towards the tree trunk, where 
they hide in crevices in the bark. It is a more northern or alpine species than 
C. boerneri although a few specimens of C. laricis have been collected with the more 
abundant C. boerneri in Southern Britain and a few specimens of C. boerneri have 
been collected among the more abundant C. laricis in the Dolomites near Cortina. 
Inouye (1962 : 147-151) describes both fundatrices and sexuales from Japan. 

The eighth abdominal tergite of Cinara laricis usually bears 15-18 hairs dorsally 
but there may also be a ventro-lateral group of 1-5 hairs on either side bringing the 
total for the tergite up to as many as 25. 

The copious honey dew crystalizes to form the 'Larchenmanne' which is important 
to bee-keepers in central Europe. 

Lachnus laricis Hartig, 1839 was not described in sufficient detail to make it 
certain that it should be placed in Cinara, let alone be applied to any particular 
species. Hottes & Frison (1931 : 155-156) place laricis Walker as a synonym of 



i 4 8 V. F. EASTOP 

laricis Hartig but from their key it seems likely that they did not have Walker's 
species. Hottes (1953 : 158) discusses the identity of Cinara laricifex (Fitch). 
Shinji (1941 : 244-248) had C. laricicolus (Matsumura) according to Inouye, 1962 : 155. 
Cinara laricifoliae (Wilson, 1915) from Larix occidentalis in North America 
is similar to C. laricis but bears rather shorter hairs on the body and appendages 
and five sub-apical setae on the processus terminalis, according to a single aptera 
vivipara kindly provided by Mr G. A. Bradley. A single ovipara of C. lyallii 
Bradley, 1956 (leg. G. A. Bradley) from Larix lyallii in Alberta is similar to C. 
laricifoliae except for the smaller siphuncular cones and may only be a form of 
C. laricifoliae. Cinara spiculosa Bradley, 1956, from Larix laricina in Labrador, 
New Brunswick and Saskatchewan is also similar to C. laricis but is usually smaller 
and the body bears bifurcate dorsal hairs. 

Cinara pectinatae (Nordlinger) 
(Text -figs 26 & 27) 

[Aphis piceae Panzer; Nordlinger, H., 1863 : 133-137. Misidentification.] 

Aphis pectinatae Nordlinger, A., 1880 : 63. [No types. ? Germany.] 

[Lachnus piceae (F.) Altum, 1880 : 352-353. Misidentification.] 

Lachnus pichtae Mordwilko, 1895a : 84, 85, 94, 96, 103. [Types unknown. Germany: Boh- 

men, Carlsbad, Abies pectinata, 1893 (M . Pawlowa)] ; 18956 : 104-105; Cholodkovsky, 1898 : 

665; 1902 : 7; van der Goot, 191 5 : 403-405. 
Eulachnus macchiatii Del Guercio, 1909 : 316, 321-324. [Types unknown. Italy: extremities 

of the twigs of Abies pectinata, Pinus picea or Picea excelsa, in September & October.] 
Lachniella pichtae (Mordwilko) Jackson, 1919 : 164-165. 
Panimerus pichtae (Mordwilko) Theobald, 1929 : 140-142. 
Dilachnus pichtae (Mordwilko) Wellenstein, 1930 : 738, 748-749, 754. 
Cinara pichtae (Mordwilko) Borner, 1932 : 568; Mordwilko, 1948 : 202. 
Cinara pectinatae (Nordlinger) Braun, 1938 : 478, 485; Kloft, Kunkel & Ehrhardt, i960 : 165- 

166; Szelegiewicz, 1962 : 84; Shaposhnikov, 1964 : 522. 
Neochmosis pichtae (Mordwilko) Kloet & Hincks, 1945 : 72. 

Buchneria pectinatae (Nordlinger) Borner, 1952 : 41; Pasek, 1954 : 263-266; Heinze, 1962 : 160. 
Cinara {Buchneria) pichtae (Mordwilko) Bodenheimer & Swirski, 1957 : 2 4^- 

Material studied. 

England: Surrey, Wisley, Abies pindrow brevifolia, 14. vi. 1957, 8 apterae, 1 alata; 
6.viii.i953, 2 apt.; 2.ix.i954, 8 apt. (/. P. Doncaster). Cambridge, University 
botanical gdns, Abies veitchii, 10-12. x. 1950, 5 oviparae; 15.x. 1954, 3 ovip., 27.ix.1954, 
2 al. viviparae (H. L. G. Stroyan & colln.); 16. x. 1950, 1 ovip.; 11.ix.1951, 7 apt. 
viviparae; 8.X.1951, 4 oviparae; Abies numidica, 3-24.X.1951, 1 alata vivipara, 
1 ovipara & 2 alate males (V.F.E.). Scotland : Morayshire, Rothes, Abies pectinata, 
24-27. ix. 1914, 1 apt. (D. J. Jackson). 

Czechoslovakia: Vysoka, Abies alba, 16.ix.1952, 2 apt. (V. Pasek). Nether- 
lands: Wageningen arboretum, Abies pinsapo, 21. vi. 1954, 2 apt. (D. Hille Ris 
Lambers & J. P. Doncaster) ; A. pectinata, i.x.1946, 1 alate male (D.H.R.L. & colln.); 
October 1966, 3 oviparae (D. Doon coll.), D.H.R.L. colln. Poland: Lesn. Podgorze, 
G. Swietokrzyskia, A. alba, 15.vii.1962, 3 apt. (/. Karpinski), H. Szelegiewicz 



REVIEW OF CINARA IN BRITAIN 



149 



colln. 1868. Zakopane, A, alba, 29. ix. i960, 3 oviparae (R. Bielawski coll.), H. 
Szelegiewicz colln. 1430. Turkey: Borncua-Izmir, 'pine tree', 1948, 4 apt. (Nihot- 
Iyriboz). Bolu-Aladag ormani, 1500 m, A. bornmuelleriana, 4^:1.1964, 5 apt. 
(H. Qanakcioglu). 

Host-plants. Abies alba (= pectinata), A . numidica, A.pindrow subsp. brevifolia, 
A . veitchii and occasionally A . bornmuelleriana and A . pinsapo. 

Distribution. Specimens have been seen from Czechoslovakia, England, 
Netherlands, Poland, Scotland and Turkey and have also been recorded from 
Austria, Germany, Sweden, Yugoslavia and the Ukraine. 

Biology. Apterae viviparae have only been seen in Britain during June, 
September and October. Oviparae and alate males have been collected in October 
Only four British alatae viviparae are known, one was taken in June, two in Sep- 
tember and the other in early October. C. pedinatae seems not to have been 
collected in Britain during July or August. Leonhart (1940) and Fossel (1958) 
have given accounts of the biology. Gontarski (1941 : 326-327) and Zoebelein 
(1956 : 374) have investigated the honeydew. 



Notes. 



Type-species of Buchneria Borner, 1952 



The alate males have unusually strongly pigmented abdominal sternites. Hottes 
& Essig (1954) compare Cinara pedinatae with C. alacra in its original description 
and C. fornacula Hottes is another similar American species. Todolachnus abietis 
(Matsumura, 1917) is closely related according to Hille Ris Lambers (1966c : 124). 




Figs 26 & 27. Cinara pedinatae. 26, <J genitalia, x 135. 27, aptera vivipara, right 
siphunculus and part of fifth abdominal tergite, x 87. 



150 V. F. EASTOP 

Cinara piceae (Panzer) 

Aphis piceae Panzer, 1801, no. 22 (2 pp. + plate). [No types. Germany]; Zettersteat, 
1828 : 557-558; Kaltenbach, 1843 : 141-142; Walker, 1848 : 95-96. 

[Aphis laricis Walker, 1848 : 102-103, partim (see Doncaster, 1961 : 93).] 

Lachnus grossus Kaltenbach, 1846 : 174-175. [No types. Locality no stated, presumably 
Germany: near Aachen, on fichten, Pinus abies, May & June]; Mordwilko, 1895a : 96. 

[Lachnus piceae (Walker) Buckton, 1880 : 58-59, mostly, but one specimen of Tuberolachnus 
salignus was included; Mordwilko, 1895a : 99; 18956 : 100; Cholodkovsky, 1898 : 655-656; 
1899 : 468; Schouteden, 1906 : 205-207; van der Goot, 1915 : 401-402; Mordwilko, 1929 : 
27.] 

[Lachnus longipes (Dufour) Buckton, 1881 : 59-61, partim, aptera only, the alatae are Tubero- 
lachnus salignus. Misidentification.] 

[Panimerus vanduzei (Swain) Theobald, 1929 : 154-156, partim (the Irish specimens are genuine 
vanduzei = abieticola). Misidentification.] 

Dilachnus piceae (Walker) Wellenstein, 1930 : 738, 742. 

Dilachnus grossus (Kaltenbach) Wellenstein, 1930 : 747, 750, 757. 

Cinara piceae (Panzer) Borner, 1932 : 569-570; Braun, 1932 : 479, 491; Wahlgren, 1939 : 2; 
Mordwilko, 1948 : 201; Szelegiewicz, 1962 : 84-85. 

[Cinara vanduzei (Swain) Borner, 1932 : 570; Braun, 1932 : 479; Onouye, 1937 : io 5- Mis- 
identification.] 

Cinara piceae (Walker) Mordwilko, 1933 : 159. 

Neochmosis piceae (Panzer) Hille Ris Lambers, 1935 : 63. 

[Neochmosis vanduzei (Swain) Kloet & Hincks, 1945 : 72. Misidentification.] 

Cinaropsis (Mecinaria) piceae (Panzer) Borner, 1949 : 59; 1952 : 44. 

Cinaria (Mecinaria) piceae (Panzer) Pasek, 1954 : 198-200, 201. 

Cinara (Mecinaria) grossa (Kaltenbach) Aizenberg, 1956 : 131-139. 

Mecinaria piceae (Panzer) Heinze, 1962 : 166, 167, 168. 

Cinara grossa (Kaltenbach) Shaposhnikov, 1964 : 521; Pintera, 1966 : 314-316; Inouye, 
1970 : 69. 

Cinara piceae var. pasheki Szelegiewicz, 1962 : 85. [Types. Polish Academy of Science, 
Institute of Zoology: Czechoslovakia & Poland.] 

Material studied. 

England: London, South Kensington, flying over roof of BMNH, 6.V.1961, 
1 alata (/. F. Perkins). Grounds of Buckingham Palace, light trap, 25-26. vi. 1964, 
1 al. (/. D. Bradley). Kennington, Picea pedinata, June 1901, 1 aptera, 1 al. 
(F. V. Theobald). Hants, Alresford, spruce, 28.xii (prior to 1881), 9 eggs (/. Ander- 
son), Buckton colln. 339 & 340. Kent, Wye, Picea excelsa, y.vi.ign, 1 al. (F.V.T.). 
Surrey, Wisley, Picea sitchensis, 16.V.1938, 5 fundatrices; 2.vii.i938, 3 alatoid 
nymphs (F. Fox-Wilson). Wrecclesham, P. excelsa, 10. vi. 1963, 1 al. (C. I. Carter). 
Kew Gardens, oriental spruce, August 1926, 2 apt. (E. V. Laing). Banstead, 
Picea sp., October 1954, 1 apt. vivipara, 3 oviparae (B. M. Gerard). Bucks, Slough, 
spruce, October 1963, 1 apt. vivip., 2 oviparae (Royal Horticultural Society). High 
Wycombe, October 1968, 3 oviparae (/. Perry). Berks, Reading, 15.V.1961, 5 larvae, 
(Reading Museum). Newbury, Picea sitchensis, 9.VL1952, 7 al. (W. D. Empson). 
Oxon, Henley-on-Thames, P. pungens var. glauca, 24.lv. 1961, 20 fundatrices (T. 
Barnard), via A. C. Jermy. Gloucs, Cowley Manor, June 1959, 1 al. (R. S. George). 
Devon, Scots pine, June 1937, 1 apt., 3 al. & alatoid nymph (L. N. Staniland). 
Dorset, Broadstone, P. spinulosa, 18. ix. 1952, 7 apt. (C. Carter). Derbys, Alport, 



REVIEW OF CINARA IN BRITAIN 151 

Hope Forest, P. sitchensis, 19.vii.1965, 1 al. (C. /. Carter). Cambridge, Picea sp., 
26.x. 1950, 47 oviparae (H. L. G. Stroyan). Cumberland, Penrith, spruce, i.x.1913, 

1 apt. vivip., 1 ovipara (F. R. Markham). Keswick, silver fir, 20.vi.1961, 1 al. 
(P. Becker). Westmorland, without further data, 3 al. & 2 alatoid nymphs (G. B. 
Piffard). Yorks, Allerston Forest, P. excelsa, 13.vii.1949, 2 al. (H. S. Hanson). 
Also in the BMNH collection are 9 alatae without data on Walker slides 638-646, 

2 alatae from spruce collected by G. Saunders on G. B. Buckton slide 336 and 1 ovi- 
para, 29. xi. — , Buckton slide 341, on which Buckton's interpretation of longipes 
Dufour is based. Scotland: Lochgoilhead, July 1922, 1 al. [A. Cuthbertson) . 
Novar, spruce, 31.x. 1923, 1 ovipara [A. S. Watt). S. Queensferry, Hopetown, 
sitka spruce, 28.ix.1931, 1 apt. vivip., 4 oviparae (G. C. R. McLaggan). Aberdeen, 
Dee Valley, spruce, 29. vi. 1965, 1 al. (M. Crooke). Eire: Dublin, Picea sitchensis, 
October 1959, 4 oviparae. 

Austria: Saalbaach, at light, 29.vii.1962, 1 al. ; 'Yellow Composite Stars', 
30.vii.1962, 2 al. (M.C.); Spider's web, 30.vii.1962, 1 al. (M. Clifton). Gross 
Glochner Pass, on pine fence, 24.vii.1967, 6 al. (H. J. Banks). Czechoslovakia: 
without further data, 1 al., F. V. Theobald colln. Banska Stiavnika, Picea excelsa, 
7.V.1950, 1 fundatrix; P. orientalis, 13. v. 1950, 1 fundatrix; 23.viii.1950, 3 apt. 
Gelnica, P. excelsa, 18.vi.1952, 1 apt. Zilina, P. excelsa, 10.viii.1949, 2 apt. (V. 
Pasek). One alata collected prior to 1933 by D. Aubertin, without further data. 
France: Savoy, Glacier des Evettes, 2600 m, July 1929, 2 al. (P. Vayssiere). 
Italy: Dolomites, Mt Sella, 8-9000', 8.viii.i962, 7 al. (0. W. Richards). Nether- 
lands: Bennekom, P. excelsa, 25. ix. 1946, 3 apt. viviparae, 1 ovipara; Ede, P. 
sitchensis, 28. v. 1957, 1 fundatrix or second generation aptera (D. Hille Ris Lambers 
& colln.). 

Host-plants. Picea species, mostly of the Eupicea group; seen from Picea 
abies (= excelsa, rubra A. Dietr. nee Du Roi) including the variety fennica and more 
rarely P. glauca (= alba, canadensis), P. orientalis and also recorded from Picea 
obovata, P. pungens and P. schrenkiana (= tianschanica) . Also seen from Picea 
sitchensis of the Cascita group and P. spinulosa of the Omorika group. 

Distribution. Specimens have been seen from Austria, Czechoslovakia, Eire, 
England, France, Italy, Netherlands, Scotland and are also recorded from Belgium 
((Schouteden, 1906 : 203), Finland, Germany, Norway, Poland, Sweden, Switzerland, 
U.S.S.R., Estonia, Latvia (Zirnits, 1927 : 251), Ukraine, Kazakhstan and Japan 
(Inouye, 1970 : 69). The records from Spitzbergen are based on wrongly identified 
specimens of C. abieticola (see p. 125). 

Biology. In Southern England adult fundatrices are present by late April 
or early May and apterae viviparae have been collected in June and from August 
to October: alatae viviparae occur in May and June. Alatae are more abundant 
than apterae during June. In Derbyshire, Yorkshire and Scotland alatae occur 
during June and July. Oviparae occur in September and are the commonest 
form found during October. Males have not been seen. According to Pintera 
(1966 : 316) the eggs hatch rather late and the colonies feed first on the older branches 
and trunk and move down to the roots in the summer. Attended by ants, especially 



152 V. F. EASTOP 

Formica and Camponotus spp. Inouye (1970 : 69) gives an account of the biology 
in Japan. Cinara piceae is a large aphid which sometimes occurs in large numbers 
in parks and gardens during spring and early summer and in the autumn. It may 
cause damage to the twigs and Nuorteva (1957 : 35-36) has investigated the saliva. 
Gontarski (1941 : 321-322) and Zoebelin (1956 : 373-374) have studied the honeydew. 

Notes. 

Type-species of Mecinaria, Borner, 1949 

From Panzer's (1801) account of Aphis piceae, 'Die Blattlaus auf der Weistanne' 
it seems that he had seen both C. piceae (= grossa) from Picea and C. abieticola 
from Abies, and considered them to be the same species. He coined his scientific 
name from the host of the spruce-feeding aphid and his common name from the 
host plant of the fir-feeding aphid. His description and illustration however applies 
to the Picea-ieeding species, the reddish hind tibiae with dark apices being typical 
for the Picea-ieeding aphid and excluding C. abieticola, which has entirely dark 
hind tibiae. Panzer used the name Pinus picea for spruce, Picea abies (= excelsa) 
and used the name Pinus Abies for fir, Abies alba [== pectinata). Kaltenbach used 
Pinus Abies for Picea abies, spruce. The use of the name Picea abies for both the 
common spruce and a widely planted fir has contributed to the confusion in aphid 
literature. 

The Picea-ieeding aphid Cinara piceae is distinguished from the Abies- and 
Cedrus-ieeding species C. abieticola by 10 of the 31 characters given in Table 3 
(page 173) but C. piceae is similar in many respects to the North American C. curvipes 
(Patch) which also lives on Abies and Cedrus. 

The oviparae resemble the viviparae in the absence of pseudosensoria on the hind 
tibiae. Cinara piceae is also unusual in that while oviparae are common, males 
are unknown. The pseudosensoria on the hind tibiae of most oviparae are thought 
to emit pheromones (Pettersson, 1970 : 63-73). The fundatrices also resemble 
the later generations of apterae viviparae and are not more densely hairy as they 
are in other species of Cinara. The unusual similarity between fundatrices and 
apterae viviparae may indicate that Picea has only recently been acquired as a 
host plant. Cinara piceae is very different from the other European species of 
Cinara feeding on Picea. The fundatrices of C. piceae may be recognized by the 
processus terminalis being only 9-22 % of the length of the sixth antennal segment 
and the base of the sixth antennal segment is 0-55-0-66 times as long as the fourth 
rostral segment, while in the later generations of apterae viviparae the processus 
terminalis is 18-28 % of the length of the sixth antennal segment and the base of 
the sixth antennal segment is 0-42-0-54 as long as the fourth rostral segment. 

Cinara pilicornis (Hartig) 

(Text-figs 28-31) 

[? Aphis pineti F. ; Zetterstedt, 1828 : 558-559 (see Wahlgren, 1939 : 2). Misidentification.] 
Aphis pilicornis Hartig, 1841 : 369. [Types unknown. Locality not stated, presumably 
Germany.] 
? Lachnus pinicola Kaltenbach, 1843 : 154-155. [Types unknown. Locality not stated, 



REVIEW OF CINARA IN BRITAIN 153 

presumably Germany: on spruce, April to August]; Schouteden, 1906 : 207. 
Aphis abietis Walker, 1848 : 100. [Lectotype, BMNH, England.] 
Lachnus hyalinus Koch, 1856 : 238-240. [No types. Locality not stated, presumably 

Germany: on spruce, June, apterae and alatae]; Mordwilko, 18956 : 106-107; Cholod- 

kovsky, 1896a : 146-148; 1898 : 663-665; ? Patch, 1912 : 165-167; van der Goot, 1915 : 

394-396. 
Lachnus macrocephalus Buckton, 1881 : 48-50. [Type, BMNH: England.] 
Lachnus flavus Mordwilko, 1895a : 94, 101-102; 18956 : 133-134. [Types unknown. With- 
out locality, presumably Poland]; Cholodkovsky, 1898 : 659. 
[? Lachnus pinicola Kaltenbach; Mordwilko, 1895a : 100-101. Misidentification.] 
? Lachnus piceicohts Cholodkovsky, 1896a : 146, 148-150. [Types unknown. U.S.S.R., near 

Leningrad & Estonia, on spruce, 1895 & 1896]; 1898 : 402-403; 659-662; 1902 : 7. 
[? Lachnus pinicolus Kaltenbach; Del Guercio, 1900 : 108. Misidentification.] 
Lachniella hyalina (Koch) Del Guercio, 1909 : 303-305. 
Lachnus abietis (Walker) Wilson & Vickery, 191 8 : 27. 

[? Lachniella pinicola (Kaltenbach) Jackson, 1919 : 165. Misidentification.] 
Dilachnus hyalinus (Koch) Swain, 1919 : 213. 

[Panimerus pinicola (Kaltenbach) Theobald, 1929 : 129-131. Misidentification.] 
Panimerus pinihabitans (Mordwilko) Theobald, 1929 : 132-135. Misidentification.] 
Panimerus hyalinus (Koch) Theobald, 1929 : 152-153. 
Neochmosis abietis (Walker) Theobald, 1929 : 352-354. 

[? Cinara pinicola (Kaltenbach) Hottes & Frison, 1931 : 156-157. Misidentification.] 
[Cinara pinicola (Kaltenbach); Borner, 1932 : 568; Braun, 482. Misidentification.] 
? Cinara nopporensis Inouye, 1937 : IOO_I 37- [Types in Entomological Institute of Hokkaido 

Imperial University. Japan: Hokkaido, Picea glehni, May, June & September, 1936.] 
[Cinaropsis pinicola (Kaltenbach) Borner, 1939 : 76. Misidentification.] 
? Cinara hyalina (Koch) Blanchard, 1939 : 864-865. 
? Cinara piceicola (Cholodkovsky) Palmer, 1945 : 447-448; 1952 : 39-40 (see Hottes, 1952 : 39- 

40). 
Neochmosis hyalinus (Koch) Kloet & Hincks, 1945 : 72. 
Cinaropsis pilicornis (Hartig) Borner, 1952 : 43; Heinze, 1962 : 170-171. 
Cinara (Cinaropsis) pilicornis (Hartig) Pasek, 1954 : 207-210; Doncaster, 1961 : 13-15; 

Inouye, 1970 : 80-81. 
Cinara pilicornis (Hartig) Hottes, 1955 : 76-77; Kloft, Kunkel & Erhardt, i960 : 166; 

Szelegiewicz, 1962 : 25; Shaposhnikov, 1964 : 522; Pintera, 1966 : 304-307. 
[Cinara (Cinaropsis) pinicola (Kaltenbach) Canakcioglu, 1966 : 139. Misidentification.] 

Material studied. 

England: Francis Walker slides 663-666, each labelled only 'pint L.' and each 
bearing a single alata vivipara. F. Walker slide no. 4 labelled 'abietis only and 
bearing a single alate male. Single alata vivipara labelled only '20. vi. 1949 [Hanson). 
London, Southgate, spruce fir, 3.vii.i847, 1 alata vivipara, 1 alate male (F. Walker 
7 & 7a); 24.vii.1847, 2 al. <$<S {F.W. 1167); 16.viii.1847, 2 oviparae (F. Walker 370). 
Surrey, Wisley, young spruce, i.vii.1958, 3 alatae viviparae; spruce, May 1957, 3 al. 
(P. Becker). Picea sitchensis, 14. vi. 1957, 5 al. (/. P. Doncaster); 'Abies pindrow 
brevifolia , 14. vi. 1957, 1 al. (J.P.D.). Farnham Forest Res. Stn, Picea sitchensis, 
29. v. 1957, 1 al. (/. H. Styles). Kew Gardens, P. likiangensis, 21.V.1960, 5 apterae 
viviparae; 16.vi.1961, 3 al.; P. asperata, n.v.1961, 4 apt.; P. breweriana, 16.vi.1961, 
2 al.; P. obovata, 16.vi.1961, 2 apt., 3 al. (V.F.E.); Picea excelsa, 18. vi. 1920, 1 al. 
[F. Laing). Fernhurst, P. sitchensis, 11.vii.1967, 1 apt., 1 al. ; Middlesex, Moor Park, 
Picea, 15.vii.1962, 1 al. (V.F.E.). Herts, Whetstone, m/v light, 5.^.1961, 1 al. 



154 



V. F. EASTOP 



(P. H. Ward). Harpenden, Rothamsted, P. excelsa, 16.vi.1920, 1 apt., 2 al. (F. V. 
Theobald). Beds, Streatley, 'scots pine', g.vi.1913, 9 apt. (F.V.T.). Bucks, Slough, 
spruce, 1937, 1 apt., 3 al. (A. Downes). Berks, Reading, Picea, j.vi.ig$o, 2 al. 
(V.F.E.). Kingsmere, P. excelsa, 12.vi.1949, 5 apt., 8 al. (V.F.E. & A. Mills). 




Figs 28-31. Cinara pilicornis, aptera vivipara. 28, x 30. 29, hind tarsus, x 87. 
30, antenna, x 87. 31, sixth antennal segment, x 87. 



REVIEW OF CINARA IN BRITAIN 155 

Hants, Bramshot, 'Abies' excelsa, 26. vii. [prior to 1881], 1 ovipara, 2 alate males 
(G. B. Buckton (slide 274, type-series of macrocephalus)) . Headley, Picea excelsa, 
29. v. 1950, 4 apt. (V.F.E.). Sussex, Ashdown Forest, Picea, 5.vii.i96i, 1 al. (H. K. 
Airy Shaw). Kent, Bromley, Picea abies, 5. vii. 1965, 2 al. (H. C. Dale). Wye, 
spruce, 5.vii.i9i3, immature including 1 alatoid nymph; 2.VL1914, 5 al. ; 10.viii.1920, 
2 al. (F.V.T.). Vagrant alata, 14. vii. 1927 (F.V.T.). Cambridge, University botanic 
gdn, Picea likiangensis, 12. vi. 1952, 10 al.; P. asperata, 11. ix. 1952, 1 apt. vivipara, 

1 ovipara {V.F.E.). Picea sp., 26.x. 1950, 3 oviparae (H. L. G. Stroyan & colln.). 
Staffs, Burton-on-Trent, Picea sitchensis (as Abies menziesii), 5. vii. 1847, x a ^- (F- 
Walker, 369). Cumberland, Great Salkeld, spruce, 2-3.vi.1911, 1 apt., 4 al.; 
4.V.1913, immature; 26.vi.1914, 1 apt., 2 al. Penrith, spruce, 3.vi.i9ii, 6 al. 
(F. V. Theobald). Ennerdale, P. sitchensis, 3. vii. 1945, 2 al. (H. S. Hanson). Scot- 
land: Aberdeen, Seaton's nursery, 'on broom', July 1926, 5 apterae viviparae, 

2 of them ovipariform and 2 alatae viviparae (E. V. Laing). Sutherland, R. Shin, 
2 m. west of Inveran, Picea abies, 9.vii.i96i, 3 al. (/. P. Doncaster). Caithness, 
Rumster Forest, P. sitchensis, 4 apt., 1 alatoid nymph (C. /. Carter). Wales: 
Anglesey, Wern Forest, Picea, 4.VL1963, 6 apt. (V.F.E.). Ireland: Rathdrum, 
Pinus sylvestris, 22.vi.1912, 1 al.; spruce, 22.V.1913, 2 apt. (F. V. Theobald). 

Austria: Gross Glochner Pass, pine fence, 24. vii. 1967, 1 al. (H. J. Banks). 
Carinthia, Ossiach, Picea, 8.viii.i966, 1 apt. {V.F.E.). Czechoslovakia: Gelnica, 
P. excelsa, 17. vi. 1952, 3 apt.; Palana Mountain, P. excelsa, 22.vi.1951, 2 apt., 1 al. 
(V. Pasek). Germany: Zucht von Schleipitz, P. excelsa, 5.XL1940, 1 ovipara 
(K. Heinze). Gratrath, P. excelsa, 16. vii. 1952, 1 apt. (H. Schmutterer coll., K. 
Heinze leg.). Rottbitze nr Bonn, Picea seedling, 18.viii.1966, 9 apt. (V.F.E.). 
Norway: Espeland, Fana, trapped, 28. vi. 1954, 1 al. (H. Tambs-Lyche). Sweden: 
Trey-toa, Picea pungens, 16. vii. 1949, 1 al. (F. Ossiannilsson). Turkey: Istanbul, 
Bahcekoy, no m, P. abies, 5.VL1964, 8 apt., 8 al.; Artvim, Kuruki'irun, 1940 m, 
P. orientalis, 8.viii.i964, 4 apt., 1 al. (H. Canakcioglu). 

Australia: N.S.W., Bilpin, Picea abies, 10.viii.1967, 1 apt., 1 al. (M. Casimir 
coll.), M. Carver leg. 

Host-plants. Picea species of the Eupicea and Cascita groups ; specimens have 
been seen from Picea abies (= excelsa, rubra A. Dietr. nee Du Roi), P. asperata, 
P. obovata, P. orientalis and are also recorded from P. glauca (= alba, canadensis), 
P. koyamai (= koraiensis), P. pungens, P. rubens (= rubra Du Roi), P. schrenkiana 
(= tianschanica) of the Eupicea group; seen from P. likiangensis, P. sitchensis and 
also recorded from P. engelmanii of the Cascita group and one sample has also been 
seen from Picea breweriana of the Omorika group. 

Distribution. Specimens have been seen from Austria, Czechoslovakia, England, 
Germany, Ireland, Norway, Scotland, Sweden, Turkey, Wales and New South 
Wales, Australia. Cinara pilicornis is also recorded from France, Hungary, Iceland, 
Japan, Netherlands, Poland, Roumania, Yugoslavia, U.S.S.R., Latvia, Lithuania, 
Estonia, Georgia, Ukraine, Khazakhstan, Caucasus and is said to have been intro- 
duced to America (Patch, 1912; Burnham, 1938; Archibald, 1958 : 106). American 
specimens resembling pilicornis seen by the present author appear to belong to 



156 V. F. EASTOP 

related American species such as braggii. Blanchard's (1939 : 864-865) description 
of the single alata he collected on Abies and identified with C. hyalina fits C. pilicornis 
but as there are several similar North American species, the Argentinian record 
cannot be accepted with certainty. Gomez Menor (1962 : 382-386) applies the 
name pilicornis to what appears to be a short-haired member of the pint group. 
Gomez Menor's (1934 : 1372) record of Cinara pinicola from Dominica was probably 
not based on pilicornis either. 

Biology. Apterae viviparae can be collected from May to July and one specimen 
has been seen from September. Alatae viviparae occur from May to August and 
sexuales from July to September. A much higher proportion of viviparae are winged 
than in other British Cinara and the sexuales start to occur earlier in the year than 
usual in Cinara. Cholodkovsky (1895 : 659-662) gives an account of the biology 
of what he calls Lachnus piceicola near Leningrad and in Estonia, saying that 
sexuales occur from mid June onwards. Pintera (1966 : 304) lists papers dealing 
with the biology of C. pilicornis which in botanical gardens is only casually attended 
by Lasius niger. Borner & Franz (1956 : 308) attribute the wide geographical 
distribution of Cinara pilicornis to its comparative independence of ants. Other 
authors state that there is a close association with ants. These discrepancies may be 
due to confusion with Cinara stroyani (= piceicola auct.). Kurir (1964 : 139-157) 
gives an account of an outbreak in Austria of C. pilicornis and of its natural enemies. 

Cinara pinea (Mordwilko) 

(Text-figs 32-33) 

[? Aphis pini L., 1758 : 453 partim.] 
? Lachnus pineti Hartig, 1839 : 645. 
[? Aphis pilosa Zetterstedt, 1840 : 311 (see Wahlgren, 1939 : 2-3).] 
[Aphis pini L., Walker, 1848 : 96-98.] 
[Aphis laricis Walker, 1848 : 102-103 partim, the alatae and perhaps var. 3 (see Doncaster, 

1961 : 93).] 
[Lachnus pineti (F.) Koch, 1855 : 230-232; Cholodkovsky, 1892 : 74.] 
[Lachnus pini (L.) ; Buckton, 1881 : 50-51; Weed, 1890 : 118.] 
Lachnus pineus Mordwilko, 1895a : 75, 76, 77, 80, 82, 94, 100. [Types unknown. Without 

locality, presumably Poland.] 1895b : 102, 126—130. 
[Lachnus pineti (Koch); Cholodkovsky, 1898 : 635-638; Schouteden, 1906 : 207; van der 

Goot, 191 5 : 405-408.] 
[Eulachnus pineti (F.) Del Guercio, 1909 : 334-337-] 
[? Lachnus pini (L.); Weed, 1890 : 118; Patch, 1912 : 168-169.] 
[? Lachniella pini (L.) Jackson, 1919 : 165.] 

[Dilachnus taeniatus (Koch) Swain, 1921 : 228-229. Misidentification.] 
[Panimerus pini (L.) Theobald, 1929 : 145-147.] 
[Dilachnus pineti (Koch) Wellenstein, 1930 : 751-752.] 
(Cinara pini (L.) Hottes & Frison, 1931 : 156; Borner, 1939 : 76; 1952 : 41; Pasek, 1954 I 

134-138.] 
Cinara pinea (Mordwilko) Borner, 1932 : 569; Mordwilko, 1933 : 159; Braun, 1938 : 478, 

492; Knechtel & Manolache, 1943 : 217-219; Palmer, 1952 : 40; Heinze, 1962 : 158-159; 

Szelegiewicz, 1962 : 85-86; Shaposhnikov, 1964 : 523; Pintera, 1966 : 285-286. 
[? Cinara pinicola (Kaltenbach) ; Silvestri, 1934 : 420-421. Misidentification.] 
Cinara (Cinarella) pinea (Mordwilko) Hille Ris Lambers, 1948 : 275. 



REVIEW OF CINARA IN BRITAIN 



157 



Material studied. 

England: without further data, 2 apterae, 2 alatae, F. Walker 659-662. Kent, 
Hothfield, Pinus sylvestris, July 1925, 3 apt. ; August 1925, 1 apt. Wye, spruce, 
27.V.1913, 7 apt. & 1 alatoid nymph (F. V. Theobald). [The correct data for this 
sample is probably 'Pinus sylvestris, 5.V.1913': see data for Cinara costata from 
Wye, May 1913.] Brookland, pine, 14.V.1961, 2 apt., 1 al. (G. M. Day). Keston, 
P. sylvestris, 11. v. 1965, 6 apt.; Hosey Hill, P. sylvestris, 6.vii.i964, 2 apt. (H. C. 
Dale). London, Southgate, P. sylvestris, 2-5. vi. 1847, 3 apt., 1 al. ; 25-26. vi. 1847, 
1 apt., 1 al. ; 3.vii.i947, 1 apt., 1 al. ; 25.viii.1847, 1 apt., F. Walker 668-673. Surrey, 
P. sylvestris, 26.vii.1927, 2 apt.; Woking, Pinus, 20.V.1912, 6 apt., many alatoid 
nymphs; P. sylvestris, 19.iv.1913, 3 apt.; 6-20.V.1913, 6 apt., 2 al. & 1 alatoid nymph ; 
19.vi.1913, 1 apt. (F. V. Theobald). Woking, P. sylvestris, 15. v. 1920, 2 apt., 1 al. 
(F. Laing). Send, P. sylvestris, 9.vii.i956, 1 al. (D. J. Williams). Wisley Common, 
? Pinus, 3.V.1965, 2 apt. (P. S. Broomfield); pine, 23. v. 1965, 1 apt. (A. Stubbs). 
Weybridge, under pine with ant, 1 apt. (F. V. Theobald). Kew Gardens, Picea 




Figs 32 & 33. Cinara pinea, right siphunculus and part of fifth abdominal tergite, x 87. 

32, alata. 33, aptera. 



i 5 8 V. F. EASTOP 

excelsa, 18.vi.1920, 1 al. (F. Laing); rose bushes, 1-26. vi. 1923, 2 al. (C. L. Withy- 
combe). Oxshott, pine, 26. vi. 1923, 1 al., 13. vi. 1926, 2 apt. (0. W. Richards). 
Wisley, R.H.S. gdns, P. sylvestris, 14. vi. 1957, 1 apt., 4 al. (/. P. Doncaster) ; 
22.vi.1961, 1 al. (P. Becker). Byfleet, Sheerwater Woods, P. sylvestris, 22. iv. 1949, 

3 fundatrices (H. L. G. Stroyan & colln.). Berks, Reading, P. sylvestris, 18. vi. 1948, 

1 apt.; 30. v. 1949, 2 apt. (V.F.E.). Silwood Park, 40' suction trap, 4.x. 1968, 1 alate 
male (Forestry Commission colln.). Hants, Liphook, P. sylvestris, 26.V.1918, 1 al. 
(W. C. Crawley). Alice Holt Lodge, P. sylvestris, 24. v. 1966, 2 apt., 2 al. (C. I. 
Carter); 14. vi. 1967, 2 apt., 2 al. (H. C. Dale). New Forest, nr Brook, 10.vii.1962, 

4 apt., 1 al. (/. Grant, G. Day, P. H. Ward). Herts, Ayot St Lawrence, Pinus 
contorta, 25. v. 1946, 4 apt., 2 al. (J.P.D.). Whetstone, m/v light, 3-25.vi.1961, 

2 al. (P. H. Ward). Cambridge, University Botanic gdn, P. sylvestris, 10. x- 
17.xi.1950, 3 oviparae; 21.V.1952, 3 apt.; 20.viii.1951, 1 apt. (V.F.E.). Suffolk, 
Barton Mills, P. sylvestris, 24.iii.1948, 6 fundatrices (H. L. G. Stroyan & colln.). 
Walberswick, P. sylvestris, 24.viii.1952, 3 apt. (J.P.D.). Brooms Barn, nr Bury 
St Edmunds, 40' trap, i.vii.1969 (N. R. Maslen, Forestry Commission colln.). 
Derbys., Wensley, P. sylvestris, 28. vi. 1946, 7 apt. (J.P.D.). Staffs, Cannock, 
corsican pine, June 1961, 1 apt.; Salop, Whixall Moss, P. sylvestris, 28. vi. 1969, 
1 apt. (B. R. Pitkin). Harper Adams, trapped, 3.x. 1958, 1 al. (J.P.D.). Cumber- 
land, Ennerdale, P. sylvestris, 3.vii.i945, 6 apt., 1 al. (M. S. Hanson). Eskdale, 
P. sylvestris, 4.vii.i953, 4 apt., 1 al. (J.P.D.). Westmorland, Windermere, P. 
sylvestris, 15.vi.1912, 2 apt. (F.V.T.). Scotland: P. sylvestris, August 1920, 

3 apt. (McDougall & F.V.T.). Kincardineshire, nr Aberdeen, P. sylvestris, July/ 
August 1919, 2 apt., 1 al. (F. Laing). Inverness, P. sylvestris, 12.viii.1948, 2 apt., 
1 al. (H. S. Hanson). Perth, Trossachs, juniper, 23. vi. 1932, 1 al. (W. H. T. Tarns). 
Angus, Glen Doll Lodge, Pinus ? contorta, 28. vi. 1959, 6 apt. (/. P. Doncaster). 
Aberdeenshire, Dinet, Pinus, 24. v. 1966, 1 apt., 1 al. (L. A. Mound). Fife, Dundee, 
40' trap, I7.ix-i4.x.i968, 2 alate males (Forestry Commission colln.). 

Czechoslovakia: Jakubov, Pinus sylvestris, 26. iv. 1952, 2 fundatrices; 18. v. 1950, 
3 apt. (V. Pasek). Kurdejov, P. sylvestris, 28. v. 1964, 1 al.; Sklene, P. sylvestris, 
30.V.1964, 1 apt., 1 al.; Kostelni Llota, P. sylvestris, i.vi.1964, 1 apt., 3 al. (V.F.E.). 
Germany: Berlin, Hohenstein, P. sylvestris, May 1939, 3 apt. ? fundatrices (W. 
Storopys), D.H.R.L. colln. Italy: Moden, P. sylvestris, August 1923, 1 apt. (C. 
Menozzi). Netherlands: Kootwk, P. sylvestris, 16.V.1929, 3 apt., ? fundatrices 
(D.H.R.L. & colln.). Bennekom, P. sylvestris, 12. vi. 1964, 2 apt., 2 al. (J.P.D. & 
D.H.R.L.). Norway: Lapland, Immerfoss, pine, 10.viii.1930, 2 apt. (Oxford 
University Lapland Expedition). Poland: Gourein, Grodno (Bieloviesch), P. 
sylvestris, 11.vii.1908, 4 apt., 1 intermediate, 2 al. (A. Mordwilko). Turkey: 
Istanbul, Baticekoy, P. brutia, 15.vii.1963, 1 al.; Istanbul, P. sylvestris, 6.V.1964, 
1 apt., 1 al.; 5.VL1964, 6 al.; Eskisehir-Fidanlik, P. nigra, 8.VL1964, 2 apt.; Bolu- 
Aladag, 1360 m, P. sylvestris, 3^1.1964, 5 apt., 2 al. (H. Canakcioglu). Yugo- 
slavia: Lesce pri Bledu, P. sylvestris, i.viii.1967, 1 apt.; near P. sylvestris, 29.vii- 
i.viii.1967, 4 apt. (V.F.E.). 

U.S.A.: Iowa, Ames, Pinus sylvestris, 11. vi. 1924, 3 apt. (F. C. Hottes). Utah, 



REVIEW OF CINARA IN BRITAIN 159 

Logan, P. sylvestris, 30. ix. 1939, 1 ovipara, 1 alate male (W. P. Nye coll., G. F. 
Knowlton leg.); pine, 5.VL1942, 1 apt. (E. Stoddard coll., G.F.K. leg.); Pinus nigra, 
5.x. 1937, 1 apt. viviparae, 2 oviparae, 1 alate male, (G.F.K. & F.C.H.); 9^.1928, 
2 apt. (G.F.K.); 5.x. 1940, 2 apterae viviparae, 2 oviparae (W. P. Nye coll., G.F.K. 
leg.). Foot of Mount Logan, aspen, 1938, 7 apt. (R. Nye coll., G.F.K. leg.). 
Canada: Manitoba, Winnepeg, P. sylvestris, 15. vi. 1964, 2 apt., 2 al. (A. G. Robinson). 
New Brunswick, Frederickton, P. sylvestris, 12. vi. i960, 1 al. (/. B. Adams coll., 
M. E. MacGillivray leg.). 

Host-plants. The usual host-plant is Pinus sylvestris and occasionally in dry 
areas, it occurs on P. nigra. Cinara pinea is also recorded from five other members 
of the Lariciones group, P. densiflora, P. kesiya (= khasya, insidaris), P. mugo, 
P. hamata (= sosnovskyi) and P. thunbergii. C. pinea is also recorded from Pinus 
banksiana (= divaricata) , P. contorta and P. halepensis of the Insignis group and from 
Pinus scopulorum of the Australes group. 

Distribution. Specimens of Cinara pinea have been seen from Czechoslovakia, 
England, Germany, Italy, Netherlands, Norway, Poland, Scotland, Turkey, Yugo- 
slavia, U.S.A., Iowa, Utah; Canada, Manitoba, New Brunswick and Ontario. C. 
pinea is also recorded from Austria, Belgium, Bulgaria (Tashev, 1961 : 157), 
Hungary, Portugal (Ilharco, 1968a : 119), Roumania, Sweden, Switzerland, 
U.S.S.R., Estonia, Latvia, Lithuania, Ukraine, Georgia and Eastern Siberia, with 
queried identity (Grechkin, 1962 : 707), Wales (Thomas & Jacob, 1940 : 139, as 
C. pini) and Minnesota (Oestlund, 1922 : 118). Japanese records are now thought 
(Inouye, 1970 : 65) to apply to Cinara piniformosana (Takahashi). 

Biology. Mr H. L. G. Stroyan has provided adult fundatrices collected between 
late March and late April in England. Apterae viviparae and a few alatae occur 
in May, both apterae and alatae viviparae are common in June and apterae viviparae 
are common but alatae are rarer in July. Apterae viviparae but no alatae have 
been collected in August. C. pinea has apparently not been collected in September 
in England. Oviparae and alate males occur in October, and one alata vivipara 
has been trapped in October. C. pinea lives on the young twigs and according to 
Withycombe (1923 : 532) is preyed upon by Hemerobius nitidulus. 

Notes. The alatae viviparae of Cinara pinea occur in three forms: 

Length of first segment of Total number of secondary rhinaria on both 

hind tarsus sides of body on antennal segments 

III IV -f V 
300-3101X 15-23 0-3 

255-285 [x 6-9 4-6 

2IO-240[X !5-27 4-IO 

This pattern is typical of many aphids (e.g. Rhopalosiphum maidis) in which the 
ipterae tend to be larger than the alatae and the alatae have more secondary 
"hinaria than the apterae. The number of secondary rhinaria on the third antennal 
.egment is correlated both with size and with degree of alatiformity, while the number 



i6o V. F. EASTOP 

of secondary rhinaria on the fourth and fifth antennal segments is correlated 
almost entirely with degree of alatiformity and is almost independent of size. The 
fourth and fifth antennal segments of aphids tend to be less affected by body size 
than is the length of the third antennal segment. The number of secondary 
rhinaria on the fourth and fifth antennal segments thus tend to be inversely corre- 
lated with body size. Similarly apterae with the first segment of the hind tarsi 
less than 240^ long are usually rather small, body length 3-1-4-5 mm but bear 
1-3 secondary rhinaria in total on antennal segments IV + V. Apterae viviparae 
bearing secondary rhinaria on antennal segments IV or V rarely have the first tarsal 
segment of the hind leg exceeding 270 pt long. 

As the fundatrices are similar to the later generations of apterae viviparae the 
data are pooled in Table 1 (between pp. 172-173). The fundatrices have from 
35-49 hairs on the fifth abdominal tergite between the siphunculi where the later 
generation bear 18-40 hairs. 

Dale (1969 : 270) has illustrated the siphuncular aperture of Cinara pinea. 

Dr D. Hille Ris Lambers informs me that his (1931 : 3) record of pineti from 
Pinus austriaca in Italy was based on a member of the C. pint group. Blanchard 
(1926 : 331-332, 1939 : 868-870) records Cinara pineti from Argentina but these 
records could apply to another species, possibly to C. excelsae H.R.L. 

Cinara pini (L.) 

Aphis pini L., 1758 : 453. [No types. Sweden: Pinus sylvestris.] 

Aphis nuda pini DeGeer, 1773 : 27-39. [Types unknown. Locality not stated, presumably 

Sweden.] 
Aphis pilosa Zetterstedt, 1840 : 311 partim. [Types. Entomologische Museum, Lund; 

Lappland] (see Wahlgren, 1939 : 2). 
Lachnus pini (L.) Kaltenbach, 1843 : 155-157; Koch, 1855 : 234-236. 
Aphis pinicola (Kaltenbach) Walker, 1848 : 98; 1852 : 955-956 partim, nee Kaltenbach, 

i8 4 3- 
? Lachnus pini (Kaltenbach); Mordwilko, 1895a : 98; 18956 : 98-99. 
[? Lachnus (aeniatus (Koch) Mordwilko, 18956 : 100, 124-126, partim (see Szelegiewicz, 
1962 : 2, footnote). Misidentification.] 
? Lachniella picta Del Guercio, 1909 : 293-294. [Types unknown. Italy: Monte Boni, 

Pinus sylvestris, 27. v. 1905, aptera vivipara.] 
? Eulachnus abameleki Del Guercio, 1909 : 316, 329-331. [Types unknown. Locality not 

stated, presumably Italy, at the extremities of the branches of Pinus sylvestris.'] 
? Eulachnus nudus (De Geer) Del Guercio, 1909 : 339-341. 
[Panimerus pinihabitans (Mordwilko) Theobald, 1929 : 132-135 partim. Misidentification.] 
[Cinara pineti (Fabr.) Hille Ris Lambers, 1931 : 3. Misidentification.] 

Cinara pini (L.) Borner, 1932 : 569; Braun, 1938 : 479-480; 491-492; Hille Ris Lambers, 

1948 : 274-275; Szelegiewicz, 1962 : 86; Shaposhnikov, 1964 : 523; Pintera, 1966 : 296-299. 

? Cinara guadarramae Mimeur, 1936 : 33-36. [Types, Institut Pasteur. Spain: Sierra de 

Guadarrama, 1,400 m, Pinus sylvestris, 10. ix. 1935, apterae & alatae viviparae.] 
? Cinaria montanicola Borner, 1939 : 76. [Types. Deutsches entomologisches Institut. 

Central & South Germany, Pinus montana]; 1952 : 43. 
? Cinaria setosa Borner, 1950 : 2-3. [Types. Deutsches entomologisches Institut, Austria: 
Pinus montana.'] 
Cinaria longirostris Borner, 1950 : 3. [Types. Deutsches Entomologisches Institut, 
Austria: Pinus sylvestris.] 



REVIEW OF CINARA IN BRITAIN 161 

Cinaria polyseta Pasek, 1951 m.s. (see Pintera, 1966 : 296). 

[Cinaria nuda (Mordwilko) Borner, 1952 : 42-43; Pasek, 1954 : 143-147, 149; Heinze, 1962 : 

166. Misidentification.] 
[? Cinaropsis pilicornis (Hartig) ; Gomez-Menor, 1962 : 382-386. Misidentification.] 

Material studied. 

England: Sussex, on Pinus sylvestris, Storrington, 17. v. 1928, 1 aptera, 4 alatae 
& nymphs (E. King); Forest Row, on young growth, 22. vi. 1964, 4 apt., (E. J. 
Gatt). London, Southgate, P. sylvestris, 26.vii.1848, 3 apt., 1 parasitized alata; 
4.viii.i847, 7 apt., 2 al.; 19.x. 1847, apterae viviparae & oviparae (F. Walker) 
679-683. Surrey, Alice Holt, P. sylvestris, old wood, 4.vii.i96i, 7 apt., 2 al. (C. /. 
Carver & V.F.E.). Hants, New Forest, near Lyndhurst, pine, August 1922, 2 al. 
(F. Laing). Cambridge, University botanic gdn, P. sylvestris, 14. xi. 1950, 1 ovipara 
(V.F.E.). 

Czechoslovakia: Bianska Stiavnika, P. sylvestris, 10. v. 1950, 1 fundatrix; 
9. v. 1949, 4 fundatrices; Pinus montana, 9.VL1952, 1 apt.; P. montana var. pumilio, 
17. vi. 1952, 2 apt.; Gelnica, P. sylvestris, 7.VL1952, 8 fundatrices, 5 in D.H.R.L. 
colln. Malacky, P. sylvestris, 18. v. 1950, 2 al. & nymphs. Zarnovica, P. sylvestris, 
15. ix. 1952, 2 apt. (V. PaSek). Bzendec, 29.V.1964, 1 al. {V.F.E.). France: 
Colbach, P. sylvestris, 5. v. 1965, 1 fundatrix, Laurent (D. Hille Ris Lambers colln.). 
Netherlands: Bennekom, P. sylvestris, 14. v. 1947, 4 fundatrices; 30. ix. 1946, 
2 oviparae, 3 apterous males (D. Hille Ris Lambers) ; 12. vi. 1954, 5 apt., 4al. (D.H.R.L. 
& J. P. Doncaster). Norway: Espeland, Fana, trapped, 28. vi. 1954, 2 al. (H. 
Tambs-Lyche). Russia: Petrograd, P. sylvestris, 2 apt. & nymph (A. Mordwilko), 
Turkey: Bolu Aladag, 1360 m, P. Sylvestris, 3.VH.1964, 13 apt., 1 al. [H. Canakcioglu). 

Host-plants. The usual host-plant is Pinus sylvestris and there are also records 
from P. mugo including its subspecies P. m. mughus and P. m. pumilio and from 
P. nigra (= austriaca) including P. n. maritima (= calabrica, laricio) of the Lariciones 
group. There are also records from P. banksiana (= divaricata) , P. halepensis 
and P. montana of the Insignes group, from Pinus ponderosa and the subspecies 
P. p. scopidorum of the Australes group and from P. cembra. 

Distribution. Cinara pini has been seen from Czechoslovakia, England, France, 
Netherlands, Norway, Russia (Leningrad) and Turkey and has also been recorded 
from Austria, Germany, Italy, Poland, Scotland, Sweden, Ukraine, Latvia and 
Yugoslavia. There are also a number of records from North America but as far 
as known these apply to Cinara pinea. No authentic Scottish specimens have been 
seen although C. pini may well occur in Scotland. 

Biology. English apterae and alatae viviparae have been collected in May, 
July and August and oviparae in November. Dr D. Hille Ris Lambers has provided 
Dutch and French fundatrices collected in May and Dutch oviparae and apterous 
males collected in late September. Pintera (1966 : 298) comments that 'alatae 
viviparous females may occur in the course of the whole vegetative period'. In 
July Cinara pini lives on the underside of the old branches and according to Pintera 
(1966 : 297-298) it lives on the young shoots in the spring. Wood-Baker (1951 : 



162 V. F. EASTOP 

271) records C. pini feeding around the edge of a resinous scar in Northern Italy, 
behaviour reminiscent of Tissot & Pepper's (1967 : 1-10) records of associations of 
some North American Cinara species with Pine Rust lesions. The relationship 
with ants has been discussed by Kloft (i960 : 48-49) and Borner & Franz (1956 : 
306) and the honeydew has been studied by Zoebelein (1956 : 374). 

Notes. The name Aphis pini has been applied to two different species of Cinara: 
to that here called Cinara pini (L.) and to C. pinea (Mdw.). Hottes (1930 : 186-187), 
Hille Ris Lambers (1948 : 274-275) and Stroyan (1955 : 332-333) have summarized 
the arguments. The name C. pini is used here in the sense of Aphis nuda pini De 
Geer but Mordwilko's Cinara nuda is C. escherichi Borner, which Shaposhnikov 
(1964 : 523) calls C. nuda (Mordwilko). According to specimens from Leningrad 
in the BMNH collection, Mordwilko also determined specimens of C. pini as nuda. 

Cinara pini is a member of a taxonomically difficult group of species. Summer 
apterae of C. pini have 3-8 hairs on the fifth abdominal tergite but fundatrices 
have 22-36 hairs on the fifth tergite between the siphunculi. Large specimens 
in the spring have 6-8 accessory hairs on the fourth rostral segment while the usually 
smaller summer apterae viviparae bear 8-10 accessory hairs on the fourth rostral 
segments. Specimens from Pinus sylvestris have the fourth rostral segment 140- 
i8o(x long but specimens from P. montana have the fourth rostral segment i75-2i0fx 
long and probably belong to a distinct (sub-) species, C. montanicola Borner, which 
may be a synonym of C. guadarramae Mimeur. Cinara setosa Borner may be based 
on fundatrices of this subspecies. 

The summer dwarfs of Cinara pini are similar to C. palaestinensis Hille Ris Lam- 
bers, which has even smaller siphuncular cones and usually bears relatively longer 
hairs on the third abdominal tergite and hind tibiae, and lives on Pinus brutia in 
the Mediterranean region. 

Cinara canatra Hottes & Bradley from Pinus banksiana in North America is 
similar to C. pini except that the dorsum of C. canatra is pigmented and the pro- 
cessus terminalis often bears 5 or 6 sub-apical setae as in C. escherichi. Cinara 
brevispinosa Gillette & Palmer from Pinus contorta in North America resembles 
C. pini but the longest hairs on the eighth abdominal tergite of C. brevispinosa are 
only 35-50 (i. long (55-160 \x in C. pini) and the ultimate rostral segment bears only 
4 or 5 accessory hairs. Cinara taedae Tissot from Pinus rigida in North America is 
similar to C. pini but apterae of C. taedae of body length 2-1-2-7 mm have the fourth 
rostral segment 200-220 \l long (150-180 \x in pini) and the longest hairs on the 
eighth abdominal tergite are 50-65^. long and only about twice as long as the 25-35 fx 
long hairs on the third abdominal tergite (in C. pini, 55-160 fx on the eighth abdominal 
tergite and 15-33 fx on the third abdominal tergite). Cinara pinata Hottes from 
Pinus edulis and C. thatcheri Knowlton & Smith from P. ponderosa in North America 
also resemble C. pini but their fourth rostral segments bear about 22-24 accessory 
hairs (6-10 in pini). 

Cinara pinihabitans (Mordwilko) 

? Aphis pilosa Zetterstedt, 1840 : 311 partim. [Types, Entomologische Museum, Lund. 
Lappland, see Wahlgren, 1939 : 2-3.] 



REVIEW OF CINARA IN BRITAIN 163 

[Aphis pinicola (Kaltenbach) Walker, 1852 : 955-956 partim. Misidentification.] 

Lachnus pinihabitans Mordwilko, 1895a : 75, 79, 94, 98. [Types, Polish Academy of Sciences. 

Poland: Otwock near Warsaw]; 1895b : 97-98, 118-119; Cholodkovsky, 1898 : 638-640; 

1902 : 7. 
[? Lachnus taeniatus Koch; Schouteden, 1906 : 207. Misidentification.] 
Lachniella pinihabitans (Mordwilko) Jackson, 1919 : 165. 
Dilachnus pinihabitans (Mordwilko) Swain, 192 1 : 227-228. 
Panimerus pinihabitans (Mordwilko) Theobald, 1929 : 132-135 partim, only the Southgate, 

1847 and Scottish, 1917, records; the others are based on specimens of Cinara pilicornis 

and C. costata. 
Cinara pinihabitans (Mordwilko) Borner, 1932 : 569 partim; Braun, 1938 : 479; Szelegiewicz, 

1962a : 87; 1962c : 245-249; Pintera, 1966 ; 301-302; Ilharco, 1968a : 119. 
Cinaria pinihabitans (Mordwilko) Bonier, 1950 : 2. 
[Cinaria taeniata (Koch) Borner, 1952 : 42; Pasek, 1954 : 151-153; Heinze, 1962 : 166; 

Gomez-Menor, 1962 : 373-376. Misidentification.] 

Material studied. 

England: London, Southgate, Pinus sylvestris, 4^.1847, 2 alatae and nymphs, 
F. Walker, 678. Kent, Wye, 15.vii.196g, 1 al. (N. R. Maslen) Forestry Commission 
colln. Herts, Harpenden, trap F2, 17. vi. 1944, 1 al. (/. P. Doncaster). Whetstone, 
light trap, 12. v. 1959, 1 al. (P. H. Ward). Gloucs., Sodley Ponds, Castanea sativa, 
2 al., 13. vi. 1959 (R. S. George). Hants, Liphook, P. sylvestris, 26.V.1918, 1 al. 
{W. C. Crawley). Alice Holt, suction trap, 6.VL1969, 4 al. ; 29. vi. 1969, 1 al. (N. R. 
Maslen), Forestry Commission colln. Scotland: Morayshire, Rothes, Pinus 
sylvestris, 28.ix.1917, 2 larvae (D. J. Jackson). Fife, Dundee, 40' trap, 27. vi. 1968, 
1 al., Forestry Commission colln. 

Finland: Aland, Lemland, Pinus sylvestris, 1j.vi.1966, 2 a.Tptera.e (F . Ossiannilsson). 
Germany: Serbst, P. sylvestris, 26.x. 1942, 1 ovipara, 1 alate male (K. Heinze). 
Sweden : Uppsala, Bjorklinge, P. sylvestris, 31.vii.1963, 1 apt., 1 al. (F. Ossiannilsson). 

Host-plants. Cinara pinihabitans has only been seen from Pinus sylvestris 
but there are also records from Pinus mugo subsp. mughus. 

Distribution. Specimens have been seen from England, Finland, Germany, 
Scotland and Sweden and are also recorded from Austria, Czechoslovakia, Ireland, 
Poland, Portugal, U.S.S.R., Ukraine (Mamontova, 1964 : 54) and Latvia. 

Biology. Alatae occur in traps in May and the first half of June in England 
but English apterae have not been seen, perhaps they occur high up the tree. The 
males are described as alate but little else is known about the biology. 

Notes. Cinara pinihabitans has been cited as the type-species of Neodimosis 
Toth, 1935, since it was the only species included. Neodimosis is probably a lapsus 
for Neochmosis and it is not certain which species Toth studied. The name Cinara 
taeniata (Koch) has been used for several different species including C. pinihabitans. 
Braun (1938 : 479) probably had a mixture of species under both C. pinihabitans 
and C. taeniata. Theobald's (1929 : 133-135) pinihabitans consisted mostly of 
C. pilicornis and C. costata. According to Heinze (1962 : 166) C. longirostris 
Borner is similar to, if not identical with, C. pinihabitans. Cinara abamaleki 
(Del Guercio) appears to be an Italian member of this group and could be an older 
name for C. maghrebica Mimeur. 



i6 4 V. F. EASTOP 

Cinara schimitscheki Borner 
(Text-figs 34-36) 

Cinara schimitscheki Borner, 1940 : 1. [Types, Deutsches Entomologisches Institut. Central 
Europe, ends of twigs of Pinus austriaca]; 1952 : 41; Pasek, 1954 : I 73 _I 75; Stroyan, 
x 957 : 348; Heinze, 1962 : 158; Shaposhnikov, 1964 : 523; Pintera, 1966 : 287-288. 

Cinara kosarowi Tashev, 1962 : 207-210. [Types, University of Sofia. Bulgaria: Pinus 
leucodermis , 9.viii.i96o.] 

Cinara (Cinarella) schimitscheki Borner; Canakcioglu, 1966 : 139. 









'r fcV w r frj. *' 



34 






35 

Figs 34-36. Cinara schimitscheki, aptera vivipara. 34, part of right side of fifth ab- 
dominal tergite, x 87. 35, enlargement of part of 34, x 450. 36, fourth and fifth 
rostral segments. 



review of cinara in britain 165 

Material studied. 

England: Surrey, Kew Gardens, Pinus nigra, 31. v. 1968, 2 apterae; 30. vi. 1969, 
2 apt. (V.F.E.). Cambridge, University botanic gdn, Pinus nigra var. calabrica, 
9.x. 1950, 1 apt. (H. L. G. Stroyan & colln.). 

Bulgaria: Rodopy, Pinus sp. {nigra), 31.V.1959, 2 alatae (R. Bielawski, H. 
Szelegiewicz colln. 1176). Czechoslovakia: Bianska Stiavnika, Pinus nigra, 
7.vii.i952, 1 apt. (V. Pasek). Turkey: Eskisehir, Fdanlik, Pinus nigra, 8.VL1964, 
2 apt., 1 al. (H. Canakcioglu). 

Host- PL ants. Cinara schimitscheki has been seen from Pinus nigra (= austriaca), 
from its variety P. n. maritima (= calabrica, laricio), is also recorded from the 
varieties P. n. caramatica {= pallasiana), P. n. leucodermis (as Cinara kosarowi) and 
from Pinus mugo subsp. mughus. 

Distribution. Cinara schimitscheki has been seen from Bulgaria, Czecho- 
slovakia, England, Turkey and is also recorded from Austria, Germany, Hungary 
and the U.S.S.R., Crimea. 

Biology. In May and June Cinara schimitscheki lives on the young shoots of 
Pinus nigra but according to Pasek (1954) teste Pintera (1966 : 288) it lives under 
the bark of the older branches later in the year. 

Notes. So few samples are known that it is difficult to evaluate the thickened 
hairs by which Cinara kosarowi was differentiated from schimitscheki. 

Cinara stroyani (Pasek) stat. n. 

[Lachnus piceicolus Cholodkovsky ; Schouteden, 1906 : 207; van der Goot, 1915 : 402-403. 

Misidentification.] 
[Dilachnus piceicolus (Cholodkovsky) Wellenstein, 1930 : 748, 751. Misidentification.] 
[Cinara piceicola (Cholodkovsky) Borner, 1932 : 480, 491; Braun, 1938 : 480, 491; Hottes, 

x 955 : 76-77; Stroyan, 1964 : 30-31; Pintera, 1966 : 302-304. Misidentification.] 
Cinaropsis drastichi Pasek, 1951 m.s. [see Pintera, 1966 : 3026]. 
Cinaropsis taxicola Pasek, 1951 m.s. [see Pintera, 1966 : 302]. 
Cinaropsis minor Pasek, 1951 m.s. [see Pintera, 1966 : 302]. 

{Cinaropsis cistata (Buckton) Borner, 1952 : 43; Heinze, 1964 : 171. Misidentification.] 
[Clnaria {Cinaropsis) cistata (Buckton) Pasek, 1954 : 202-207. Misidentification.] 
Cinaropsis cistata var. stroyani Pasek, 1954 : 2 °7- [Types, Slovak Academy of Science, 

Bratislava. Netherlands, leg. H. L. G. Stroyan.] 
[Cinara cistata (Buckton) Ossiannilsson, 1955 : 378; Szelegiewicz, 1962 : 81; Shaposhnikov, 

1964 : 522. Misidentification.] 

Material studied. 

England: Kent, Bromley, Picea abies, 5.^.1965, 1 alata {H. C. Dale). Surrey, 
Kew Gardens, Picea obovata, 16.vi.1961, 1 al. {V.F.E.). 

Czechoslovakia: Westerheim, Picea, 14. vi. 1932, 1 aptera, 1 alata {D. Aubertin). 
Banska Stiatnika, P. excelsa, 7. v. 1949, 2 fundatrices; 17-23.vi.1952, 5 apt.; Gelnica, 
P. excelsa, 6. v. 1952, 5 fundatrices, 2 in D. Hille Ris Lambers colln.; 26.vi.1952, 
1 apt., 1 al.; Polana Mount., P. excelsa, 9.vii.i95i, 2 apt.; Zarnovica, P. excelsa, 
15.ix.1952, 1 ovipara {V. Pasek). Praha — kosire, P. abies, 4.VL1952, 2 apt. {A. 



i66 V. F. EASTOP 

Pintera). Germany: Tschdf, P. excelsa, 29.vii.1944, 1 apt. (K. Heinze). Unteres 
Weldental-schwaiswalt, P. excelsa, 15. vi. 1944, 1 apt., 1 al. (K. Heinze). Nether- 
lands: Bennekom, Picea, 21. vi. 1949, 4 al. (D. Hille Ris Lambers & H. L. G. Stroyan 
& colln.). Poland: Bydgoszcz- Jacheice, P. excelsa, 4.viii.i956, 1 imm. apt., 1 
alatoid nymph (H. Szelegiewicz) . Sweden: Brunnby, Kullen, P. abies, 25.viii.1964, 
3 apt. (F. Ossiannilsson & colln.). 

Host-plants. Cinara stroyani has been seen from Picea abies (= excelsa, rubra 
A. Dietr. nee Du Roi) and is recorded from P. pungens, which also belongs to the 
Eupicea group. 

Distribution. Cinara stroyani has been seen from Czechoslovakia, England, 
Germany, Netherlands, Poland and is also recorded from Austria, Bulgaria, Norway 
(Stenseth & Bakke, 1968 : 238), Sweden, U.S.S.R., Estonia, Latvia (Zirnits, 1927 : 
251), Ukraine. 

Biology. In Czechoslovakia, according to Pintera (1966 : 302-304) the funda- 
trices of C. stroyani occur on the two year old shoots on the shady side of spruce 
trees and are attended by ants. Alatae occur in large numbers in May in the second 
and third generations and fly to the current years' shoots to deposit their young. 
As the summer progresses the colonies move to the older branches near the trunk 
but have not yet been discovered on the roots. According to Borner & Franz 
(1956 : 307) C. stroyani is mostly on the roots in summer. Oviparae are recorded 
from July onwards. Saemann (1966 : 380) and Kloft (i960 : 49) have also written 
on the biology and Zoebelein (1956 : 380) has studied the honeydew. 

Notes. Records of Cinara piceicola between 1915 and 1966 mostly apply to 
C. stroyani but Inouye (1938 : 80) had C. horii Inouye, 1956. Palmer's (1952 : 39- 
40) description of metatypes of C. piceicola suggest that it is a synonym of C. 
pilicornis. Continental authors suspecting this synonymy used the name cistata 
Buckton for the species. Buckton's specimen is Cinara costata from Walker's 
collection and had been correctly determined by Walker, but the '0' in costata was 
not closed and Buckton misread the name as cistata. 

According to Pintera (1964 : 304, 306) C. stroyani is very common in Central 
and Northern Europe. It is not common in England and this, together with the 
difficulty in separating stroyani from pilicornis, may indicate either an ecological 
difference between the two species or possibly that they are only forms of the same 
species. 



Cinara tujafilina (Del Guercio) 

(Text-figs 39-4 1 ) 

Lachniella tujafilina Del Guercio, 1909 : 288, 311-312. [Types unknown. Italy: nr Firenze, 
Thuja, 15. vi. 1905, aptera.] 

Lachnus thujafilinus (Del Guercio) Davidson, 1914 : 127. 

Lachniella thujafolia Theobald, 1914 : 335-336- ['Paratypes' in BMNH with data as given in 
the original description, South Africa: Transvaal, Ondersteport & Pretoria, Thuja 
orientalis, April & August, 1913, apterae viviparae. The specimen labelled 'type' in 



REVIEW OE CINARA IN BRITAIN 167 

Theobald's collection is an alata vivipara, a form not mentioned in the original description, 

collected from the Orange Free State, 27.viii.1914, a locality not mentioned in the original 

description.] 
Lachnus thujafolia (Theobald) Takahashi, 1921 : 81. 
Lachnus biotae van der Goot, 1917 : 161-163. [Types unknown. Java: Buitenzorg, 250 m, 

Biota orientalis, December, 1914, apterae & alatae viviparae.] 
Lachnus tujafilinus (Del Guercio) Swain, 1919 : 50; Zimmerman, 1948 : 63-64. 
Dilachnus thujafolia (Theobald) Hall, 1926 : 3; Okamoto & Takahashi, 1927 : 144. 
Dilachnus callitris Froggatt, 1927 : 56-58. [Types unknown. Australia: New South 

Wales, Callitris, 1921 & 1925.] 
[? Dilachnus juniperi (F.); Nevsky, 1929 : 350-351. Misidentification.] 
Cinara thujafoliae (Theobald) Takahashi, 1931 : 24; Mimeur, 1934 : 1. 
Dilachnus tujafilinus (Del Guercio) Smith, 1932 : 86. 
Panimerus thujafoliae (Theobald) Lepiney & Mimeur, 1932 : 128. 
Cinara tujafilina (Del Guercio) Borner, 1932 : 570; Boudreaux, 1948 : 98; Smith, Martorell 

& Escolar, 1963 : 52-53. 
Cinara winokae Hottes, 1934 : I_2 - [Type, USNM; U.S.A., Louisiana, Arbor Vitae, 

31.xii.1931, alata vivipara.] 
Neochmosis tujafilina (Del Guercio) Hille Ris Lambers, 1935 : 63. 
[Cinara juniperi (De Geer) ; Blanchard, 1939 : 866-868; Waterston, 1949 : 7; ? Dzhibladze, 

1958 : 293 partim. Misidentification.] 
Cupressobium tujafilinum (Del Guercio) Borner, 1952 : 44-45; Heinze, 1962 : 176. 
Cupressobium thujaphilinum (Del Guercio) Borner & Heinze, 1957 : 57. 
Cinara [Cupressobium) tujafilina (Del Guercio) Eastop, 1958 : 92. 
Cupressobium tujafilina (Del Guercio) Gomez-Menor, 1962 : 393-397. 
? Cupressobium mediterraneum Narzikulov, 1963 : 113-117. [Type, Pavlovsky Institute of 

Zoology & Parasitology. U.S.S.R., Tadzhikistan, Dushanbe, 29. v. 1958, M.N.N.] 

Material studied. 

England: Surrey, Wimbledon, Thuja, 5.VL1935, 2 apterae (C. T. Gimmingham) ; 
Wallington, cypress, June 1965, 28 apt. (Public Health Dept). Essex, Woodford, 
fir, 5.vi.i944, 4 apt. Berks, Reading, Chamaecyparis lawsoniana, 9.V.1948, 5 apt.; 
Cupressus sp., 3.^.1950, 11 apt. (V.F.E.). Hants, Alice Holt, suction trap, i.vii.1969, 
1 alata (N. R. Maslen), Forestry Commission colln. Cambridge University Botanic 
gdn, Thuja orientalis var. elegantissima, 9.V.1952, 2 apt.; Thuja sp., 15.vii.1951, 
8 apt. {V.F.E.). 

Egypt: Cairo, Thuja orientalis, February 1918, 3 apt. (A. Alfieri). Giza, Thuja, 
12.iii.1924, 2 apt., 1 al., 2 alatoid nymphs; Gezireh, Thuja, 6.IV.1924, 6 apt., 3 al. 
{W. J. Hall). Germany: Elsheim, Friedhof, Biota orientalis, 5JX.1930, 2 apt. 
(C. Borner), K. Heinze leg. Iraq: Kirur, Thuja, 16.iii.1968, 12 apt., 2 al. (H. E. 
Knopf). Italy: Rome, Juniperus communis, 25.iii.1960, 7 apt. (F. Silvestri), 
E. Tremblay leg. (2 in Inst. Ent. Agr., Portici colln.) Japan: Thuja orientalis, 
3 apt. (R. Takahashi), ex Theobald colln. Morocco: Casablanca, Thuja occidentalis, 
May 1929, 16 apt.; Rabat, Biota occidentalis, July 1929, 1 apt., 1 al. [Bouhelier, 
see Mimeur, 1933 : 1], F. V. Theobald colln. Netherlands: Wageningen arbore- 
tum, Thuja occidentalis ohlendorffi, 19. vi. 1954, 1 apt. (D. Hille Ris Lambers & J. P. 
Doncaster). Palestine: Jerusalem, Thuja orientalis, 26.V.1946, 3 apt. (E. Swirski). 
Without locality, Thuja, 20. ii. 1938, 2 apt. (S. Dauderari). Turkey: Istanbul, 
Bahcekoy, no m, T. orientalis, 5.VL1964, 21 apt., 1 al. (H. Canakcioglu). 



1 68 



V. F. EASTOP 




Figs 37-41. Cinara tujafilina, alata vivipara. 37, wings, x 27. 38, antenna, x 100. 
39, hind tarsus, x 100. 40 & 41, siphuncular cone, dorsal and lateral views, x 240. 



REVIEW OF CINARA IN BRITAIN 169 

Eritrea: Asmara, Thuja orientalis, 15.viii.1948, 3 apt., 2 al. (G. de Lotto). 
Malawi: Mt Mlanje, 6000', Widdringtonia whytei, 28.vii.1956, 6 apt., 2 al. (A. 
McCrae); June 1958, 2 apt. (G. W. Hearn). Rhodesia: Salisbury, Callitris robusta, 
19.vi.1915, 1 al., 28. ii. 1957, 2 immature (Dept. Agric); Thuja sp., 11. v. 1928, 4 im- 
mature; 21. vi. 1928, 3 apt. (W. J. Hall). Thuja orientalis, June 1930, 1 apt.; T. 
orientalis, on roots with ants, 22. v. 1965, 1 apt., 8 immature (Dept. Agric). South 
Africa: Transvaal, Onderstepoort, T. orientalis, 6.^.1913, 1 apt., many immature 
including 1 alatoid nymph, on 4 slides, paratypes of thujafolia Theobald. Pretoria, 
T. orientalis, i.viii.1913, 4 apt., many immature (/. Bedford), paratypes of thujafolia, 
mounted on 7 slides. Orange Free State, Petrusburg, Thuja, 27.viii.1914, 1 al., 
labelled 'type' of thujafolia but the alata was not described in the original description, 
nor is Orange Free State given as a locality. Ladybrand, Thuja, 12.viii.1914, 
3 apt. ; Blomfontain, Cupressus, 30.viii.1917, 2 alatoid nymphs, F. V. Theobald colln. 
Harrismith, 1-20. iii. 1927, 2 al. (R. E. Turner). 

Nepal: Kathmandu, conifer, 14.xii.1960, 2 apt. (K. C. Sharma). 

Australia: New South Wales, Dubbo, Callitris endlichi, 12.x. 1959, 3 apt. (K. M. 
Moore). Desert cypress, 24.ix.1921, 2 apt., 2 al. (W. W. Froggatt), presumably 
part of the series from which callitris Froggatt was described. Matong State 
Forest, Callitris glauca, 17.xii.1956, n apt., 2 in N.S.W. Forest Dept. colln. Buckin- 
bong S.F., C. glauca, 20.xii.1956, 4 apt., 3 in N.S.W. Forest Dept. colln. Pennant 
Hills, C. rhomboidea, 31.vii.1958, 10 apt., 1 al.; Stahom S.F., 30.vii.1959, 2 apt., 
6 al. (K. M. Moore). A.C.T., Canberra, Chamaecyparis lawsoniana, 6JX.1959, 
1 apt.; Libocedrus decurrens, 3JX.1959, 2 apt. (V.F.E.). Tasmania, Coles Bay, 
Callitris tasmanica, 26. v. 1952, 2 apt. (K. A. Pickett). Victoria, Colstream, g.ix.1949, 
6 apt. (Mrs Lithgow). Melbourne, Thuja orientalis, 29. v. 1959, 7 apt., 1 alatoid 
nymph; 23. vi. 1959, 8 apt., 2 alatoid nymphs; 3-10. vii. 1959, 20 apt., 27 al.; Thuja 
occidentalis , 30. vi. 1959, 1 apt.; T. occidentalis cultivar 'Rheingold', 30. vi. 1959, 
21 apt. (V.F.E.). 

Bermuda: Paget E., Juniperus bermudiana, 15. v. 1943, 17 apt. (/. M. Waterston). 
U.S.A.: California, Riverside, Thuja occidentalis, 8.ix.i9i6, 1 apt., 1 al. (A. F. 
Swain). Texas, Crystal Springs, Thuja, 22. ii. 1936, 4 apt., 1 alatoid nymph (M. J. 
James coll.), G. F. Knowlton leg. Utah, St George, Thuja, 22. iii. 1946, 14 apt., 
3 al.; evergreen tree, 24.iii.1951 (M. S. Burmingham coll.); Green River, arbor 
vitae, 16. v. 1958, 24 apt.; Murra, Juniperus sp., 27.x. 1964, 1 apt., 4 larvae (G. F. 
Knowlton). 

Host-plants. The usual host-plant is Thuja (Biota) orientalis but specimens 
have also been seen from the following Cupressaceae, Thuja occidentalis, Callitris 
columellaris (= glauca), C. endlichi, C. priessii (= robusta), C. rhomboidea, C. 
tasmanica, Chamaecyparis lawsoniana, Juniperus virginiana, Libocedrus decurrens 
and Widdringtonia whytei. 

Distribution. Cinara tujafilina has been seen from Egypt, England, Germany, 
Iraq, Japan, Morocco, Netherlands, Palestine, Turkey, Eritrea, Malawi, Rhodesia, 
South Africa, Nepal, Australia, Bermuda, U.S.A., California, Texas and Utah. 
C. tujafilina was originally described from Italy and is also recorded from Bulgaria 



170 V. F. EASTOP 

(Tashev, 1964 : 173), Portugal (Ilharco, 1968a : 64), Spain, Mozambique (Ilharco, 
1970 : 2-3), Puerto Rico, Florida (Tissot, 1939 : 42), Louisiana and Missouri (Kring, 
1955 = 64). 

Biology. The distribution of Cinara tujafilina is much wider in the warmer 
and drier parts of the world than most other species of Cinara. Large numbers of 
apterae occur on the undersides of the branches near the trunk in June and smaller 
numbers of apterae in May and July. Alatae have not been collected in Britain. 
According to Bray (1953 : 103-107) C. tujafilina occurs on the roots in the summer. 
According to Waterston (1949) it is attended by the ant Pheidole megacephala. 
Tissot (1939 : 42), Waterston (1949 : 7) and Weigel & Baumhofer (1948 : 19-20) 
record the damage caused by this aphid and Gomez-Menor (1963 : 397) says that 
C. tujafilina is primarily controlled by Chilocorus bipustulus in Spain. Lepiney 
& Mimeur (1932 : 28) record Scymnus subvillosus as a predator in Morocco. Froggatt 
(1923 : 163) gives an account of an outbreak of the 'Cypress Pine Aphis' in New 
South Wales in 1921 but did not formally describe the aphid and apply a latin 
binomen until 1927. 

Notes. Cinara tujafilina is more common on Thuja orientalis and Cinara cupressi 
is more common on Thuja occidentalis but Blanchard's description of Cinara juniperi 
from Thuja occidentalis in Argentina seems to apply neither to juniperi nor to 
cupressi but to C. tujafilina. Tashev (1944 : 173) on the other hand may have had 
Cinara cupressi. 

A single aptera (body length 2-5 mm) and 4 larvae collected on Juniperus excelsa, 
Turkey, Bardur Sarnie, 1300 m, 3.VU.1970 by Dr H. Canakcioglu are similar to 
C. tujafilina except that the longest hairs on the hind tibiae are only 128 pi long. 
The longest hairs on the hind tibiae of C. tujafilina are 140-240PL long and are only 
less than 155 \i in specimens less than i-8 mm long. The Turkish specimens probably 
represent a hot weather form of C. tujafilina but may be a distinct (sub-) species, 
possibly C. mediterraneum (Narzikulov) . 

HOST PLANTS OF THE BRITISH SPECIES OF CINARA 

PINACEAE 
Abies spp. 

Cinara abieticola, C. pectinatae 
Cedrus spp. 

Cinara abieticola 
Larix and Pseudolarix spp. 

Cinara boerneri, C. kochiana, C. laricis 
Picea (Eupicea) abies (= excelsa) glauca (= alba, canadensis), orientalis, pungens 

Cinara bogdanowi, C. costata, C. piceae, C. pilicornis, C. stroyani 
Picea (Cascita) engelmanii, jezoensis, likiangensis, sitchensis 

Cinara costata, C. piceae, C. pilicornis 
Picea (Omorika) breweriana, omorika, spinulosa 

Cinara costata, C. piceae 
Pinus (Insignes) banksiana (= divaricata), contorta, halepensis, montana 



REVIEW OF CINARA IN BRITAIN 171 

Cinara pinea, C. pini 
Pinus (Lariciones) mugo including mughus 

Cinara pinea, C. pini, C. pinihabitans 
Pinus (Lariciones) nigra (= austriaca) including caramatica (= pallasiana), leuco- 
dermis, maritima (= calabrica, laricio) 

Cinara acutirostris, C. brauni, C. schimitscheki and occasionally C. pinea 
Pinus (Lariciones) sylvestris including hamata (= sosnovskyi) 

Cinara escherichi, C. pinea, C. pini, C. pinihabitans 
Pinus (Australes) ponderosa including scopulorum 

Cinara pinea, C. pini 

CUPRESSACEAE 

Callitris, Chamaecyparis, Libocedrus spp. 

Cinara tujafilina 
Cupressus arizonica, torulosa 

Cinara fresai 
Cupressus goveniana 

Cinara cupressi 
Cupressus macrocarpus 

Cinara cupressi, C. fresai 
Juniperus bermudiana 

Cinara tujafilina 
Juniperus chinensis, communis, oxycanus, oxycedrus 

Cinara juniperi 
Juniperus horizontals, sabina 

Cinara fresai 
Juniperus virginiana 

Cinara cupressi 
Thuja occidentalis 

Cinara cupressi and rarely C. tujafilina 
Thuja (Biota) orientalis 

Cinara tujafilina 

ACKNOWLEDGEMENTS 

Most of the specimens studied in the course of this work are in the collection of 
the British Museum (Natural History). Dr Lena K. Ward presented the aphids 
from a survey of Juniper insects made in Southern Britain during 1968 and 1969. 
Named specimens of Cinara have been given or lent by G. A. Bradley, C. I. Carter, 
Mary Carver, J. P. Doncaster, K. M. Harris, K. Heinze, D. Hille Ris Lambers, 
F. C. Hottes, S. Huculak, M. Inouye, G. F. Knowlton, N. R. Maslen, J. O. Pepper, 
A. Pintera, J. Powell, A. G. Robinson, Louise M. Russell, H. L. G. Stroyan and 
A. N. Tissot. Text-figures 1-13, 17-18, 22-25, 2 7 and 3 2 ~36 are the work of Mrs 
J. Palmer; 28-31 of Miss B. G. Forbes-Sempill ; 20 and 37-41 of Mrs C. A. Gosney 
and 14-16, 19, 21 and 26 of Mr Arthur Smith. Figures 20 and 37-41 have appeared 
previously in the Australian Journal of Zoology and thanks are due for the permission 
to reproduce them. 



Table i 

Biometric data for the apterae viviparae of the British species of Cinara 



piceae 

fundatrix 
kochiana 
boerneri 
laricis 
acutlrostris 

fundatrix 
pint 

fundatrix 
pinthabitans 
escherichi 

fundatrix 

brauni 



Host 


u S 


c 


a> "s 


plant 


II 

Is- 


a 

m s 


1J a> 

o-§*3 


Picea 


19 


3-2-6-7 


170-630 




10 


5-4-6-7 


340-510 


Larix 


12 


5-7-6-1 


410-530 




54 


2-4-4-1 


170-300 




29 


2-5-5- 1 


230-630 


Pinus nigra 


17 


2-8-4-1 


390-710 




2 


3-3-3-5 


290-370 


P. sylvestris 


24 


2-5-4-4 


160-700 



Length in // of antennal segments 



P. nigrfl 



pinea 


P. sylvestris 


73 


schimitscheki 


P. nigra 


9 


pectinatae 


Abies 


32 


costata 


Picea 


30 


pilicornis 




43 


stroyani 




16 


fundatrix 




7 


bogdanowi 




30 


fundatrix 




2 


ableticota 


A bies 


22 


juniper i 


Juniperus 


67 


fresai 


Juniperus 
Cttpressus 


46 


cupresst 


Cupressus 
Thuja 


88 


tujafitina 


Thuja 
Callitris 


33 



3-2-4-0 310-620 

3-7-4-2 170-460 

3-2-4-3 370-640 

4-0-4-2 300-720 



2-5-3-* 



fused 



3-1-5-1 270-700 

3-3-5-1 440-720 

2-8-5-0 180-300 

2-7-3'8 330-530 

2-1-4-7 130-390 

2-1-4-2 170-580 

3-5-4-6 200-420 

2-4-5-0 240-870 

3-4-3-8 420-560 

3-8-7-8 490-990 

2-4-3-1 360-600 

2-2—4-2 200-610 

1-8-3-2 200-570 

33 1-7-3-5 130-300 



° S •£ u 2 2 

« -° % s -a " 

absent present 



„ absent 

low present 

present present 

present 
absent 
present 

small- 
absent 
absent present 

V broken 
absent present 



absent 
from V 
absent 



■sal 

absent 

very small 

very small 
large 
very small 



present 
small 



black 
patch 
large 
very small 

present 
very small 



absent 

very small 



III 

530-n 10 

820-920 

720-980 

500-720 

410-670 

530-650 

440-470 

400-700 

460-660 

620-650 

560-710 

620-670 



IV 

180-370 
230-310 
290-460 
230-320 
140-300 
180-300 
150-200 
140-260 
180-240 
25 -33o 
220-320 
240-260 



VI 



300-550 

360-420 

390-530 

280-390 
190-370 
230-330 

190-230 
190-270 

210-280 
250-2S0 
260-360 
250-270 



120-200 
170-210 
130-200 
130-190 
100-140 
130-170 
140-160 
100-180 
100-160 
120-160 
130-190 
170-180 



+ 40-81 
+ 22-51 

+ 56-88 

4- 26-52 
+ 22-35 
+ 34-49 
+ 20-27 
+ 25-52 
+ 27-46 
+ 3i-37 

4- 42-61 
+ 35-45 



530-700 190-310 250-360 170-220 + 53-95 



500-740 
630-860 
330-550 
330-470 
250-530 
340-590 
420-530 
350-740 
380-390 
580-1100 



220-360 
27O-350 
130-210 
120-200 
120-210 
130-240 
130-180 
170-350 
150-170 
280-590 



280-400 
300-380 
160-240 
180-260 
150-280 
170-300 
130-180 
240-390 
180-190 
330-630 



140-230 
120-200 
140-200 
150-210 
1 10-180 
120-180 
130-160 
120-200 
130-160 
230-290 



+ 43-71 
4- 52-70 
+ 40-63 
4- 19-35 
+ 25-45 
4- 19-44 
+ 17-30 
4- 30-60 
4- 14-22 
+ 44-90 



very small 250-370 95-180 160-280 150-230 4- 40-70 

very small 300-580 140-240 160-270 140-220 4- 40-70 

or absent 

very small 240-450 100-200 130-210 130-160 4- 25-46 

or absent 

very small 230-330 85-150 120-170 110-170 4- 18-35 

or absent 



Length 


f rostral 


segments in fi 


IV 


V 


300-370 


120-145 


300-350 


120-150 


320-420 


120-150 


150-210 


85-105 


160-210 


80-100 


210-250 


110-130 


200-2 I 


105-115 


140-220 


70-120 


150-180 


70-100 


150-160 


85-90 


210-230 


96-106 


200-210 


100-105 


250-330 


120-150 


210-290 


110-170 


290-360 


130-165 


210-270 


80-105 


190-260 


105-125 


190-250 


100-130 


210-270 


85-125 


210-250 


100-115 


240-320 


110-140 


250-260 


100-105 


270-360 


120-150 


120-170 


60-90 


160-240 


75-120 


130-180 


70—100 


140-180 


70-90 



First segment of hind tarsus in ft 



basal 


dorsal 


diameter 


length 


45-80 


51-92 


70-80 


62-77 


53-66 


62-94 


42-53 


79-112 


34-60 


57-96 


40-46 


44-53 


37-43 


40-48 


31-46 


38-57 


36-59 


40-70 


41-44 


45-67 


37-48 


49-72 


53-60 


58-64 



33-48 

47-65 
45-60 
43-58 

35-44 
31-53 
35-57 
38-49 
35-7° 
40-50 
53-81 

31-58 
35-44 

35-44 

30-45 



137-220 
86-1 I 1 

121-155 
23-35 
27-53 
33-60 
35-54 
35-76 
40-45 
20-46 

13-28 
20-34 

19-35 

19-31 



Table i (contd.) 



Hind tarsus 
segment 



ventral 

length length 



130-210 
170—200 
170-210 
150-210 
1 10-180 
100-140 
100-130 
90-130 
110-130 
120-igo 
120-150 
150-160 
150-210 
220-330 
170-220 
190-260 
80-120 
90-150 
1 10-160 
110-150 
120-180 
110-120 
130-180 
70-100 
80-120 



Hind 
tibia 

length 
mm 



360-470 
390-430 
370-450 
360-510 
270-390 
240-310 
240-250 
210-280 
270-310 
260-320 
270-320 
340-360 
320-430 
35o-53o 
320-380 
360-450 
250-330 
300-520 
340-450 
320-400 
250-380 
230-270 
3 S 0-500 
250-350 
260-350 



80-120 240-310 1 
60-100 200-280 o 



0-1-7 
8-1-6 



Length of longest hair on 
Third abdominal 

antennal Hind tergites 

segment tibia 3 8 



27-80 
40-70 
60-70 
24-70 
40-80 
7o-95 
75-9o 
25-80 
65-90 
110-150 
40-70 
60-70 
75-120 
90-210 
70-120 
45-no 
140-170 
90-150 
60-90 
70-100 
80-150 
80-110 
190-300 
140—210 
190-250 

150-220 



44-90 
55-90 
55-90 
50-70 
65-125 
90-120 
105-125 
40-95 
90-105 
170-190 
70-90 
80-100 
90-140 
120-230 
110-150 
110-190 
170-250 
130-220 
60-105 
100-135 
So-170 
130-150 
150-330 
180-240 



5-26 
9-18 
11-25 
14-33 
35-120 
40-65 
85-90 
15-35 
15-40 
140-180 
13-17 
19-30 
1 10-190 
95-210 
45-100 
30-130 
110-150 
100-160 
60-80 
90-105 
90-150 
100-120 
160-270 
140-210 
180-230 



45-130 
70-150 
90-130 
60-100 
90-180 
110-1S0 
140-160 
55-160 
130-140 
140-160 
90-120 
120-140 
120-180 
120-230 
95-130 
130-200 
120-190 
130-180 
100-150 
120-150 
120-160 
130-150 
190-360 
150-230 
190-260 



Number of 
hairs on 
second 
antennal 
segment 

9-16 

5-" 
21-32 

7-i3 

5-9 

6-8 

6-8 

5-10 

5-6 

7-8 

9-13 

9-13 

8-1 1 

5-9 

9-13 

5-8 
n-15 

9-18 
14-22 

9-16 
12-26 
15-19 
12-24 

6-13 



Number of hairs on 

sixth antennal 

segment 

sub-apical 

Base processus 

terminalis 



180-240 120-200 170-220 



6-13 



9-17 

n-16 

15-21 

6-11 

4-7 

9-12 

7-9 

8-16 

9-12 

S-io 
11-16 

19-12 
9-16 
2-8 
6-9 
5-8 

10-17 
6-14 

11-18 
6-10 

11-20 
7-10 
9-14 
7-13 
7-12 

4-7 



6-1 1 
6-8 
6-9 

4 
(3-)4 



(3)-4 



6-7 

5-7 

5-8 

(3-)4("5) 

(3-)4 

(3")4 

3-4 

(3->4 

4("5) 

(3")4 

4(-5) 

4 

3-4 

(H3(-4) 

3 



accessory 

hairs 

on 

ultimate 

rostral 

segment 

12-17 
8-12 
22-34 
5-7 
5-7 
6-9 

6 
6-10 

7 
4-6 
8-1 1 
6-7 
5-8 
4-6 



4("5) 
4-7 
6-9 
6-8 
9-13 
9 
7-13 
(3")4(-5) 
5-7 



sub- abdominal 

genital tergites 

plate 5 8 



Number of secondary 
rhinaria on antennal segments 



48-76 
47-120 
33-55 
19-56 
18-29 
18-37 
26-34 
9-36 
32-50 

28 
39-58 
52-56 
32-69 
22-52 
50-58 
30-50 
22-42 
I3-3S 
32-65 
43-56 
18-33 

27 
30-50 
19-27 
28-44 



6-17 

6-12 
18-40 
16-36 
20-42 

4-7 
18 

3-8 
23-30 
40-50 
(5-)6 
13-18 
21-36 
18-49 
26-65 
36-56 
54-92 
30-90 
40-100 
65-95 
40-90 
80-95 
70-85 
28-55 
45-65 



o(-i) 
0-2 
o 
o-3 
0-1 

0-2 
0-3 
0(-l) 

0-2 

o(-4) 



O-I 

(o-)i(- 3 ) 
o(-l) 



i-4 

o-l(-3) 



9-17 0(-l) 0-l(-2) 

7-13 o O 

4I-64 
I8-3I 
13-23 
I2-I8 
17-21 
7-18 
18-25 
20-24 
14-23 
34-49 
IO-I5 
13-26 
27-60 
23-38 
26-52 
2O-77 
H-35 
23-4° 

J9-35 
23-40 
29-43 
18-29 

17-27 



0-2 
0-1 

1 
0-2 
o-3 
o-3 

0(-2) 



0-2 
0-2 

i-3 

(O-)I 

■-4 

0-3 



1-3 

(0-)l(-2) 



O-I (-2) (0-)l(-2) 



(2-)3 



22-30 30-50 



I6-3I 



1 1 0-1 70 140-240 100-18 



19-33 50-70 J 9-26 



(0-)l(-2) 

(0-)l(-2) 



I (-2) 
1-2 

I (-3) 
I (-2) 

(0-)l(-2) 

0(-2) 



O-I (-2) 

1-2 

i (-2) 

1 (-2) 

1 

1 (-2) 

I-2(- 3 ) 

(0-)l(-2) 

I (-2) 

(O-)I 



Table 2 



; data for the alatae viviparae of the British species of Cinara 



Eg 



piceae 

hochiana 

boerneri 

laricts 

acutirostris 

pint 

pinihabitans 

escherichi 

brauni 
pinea 

schimitscheki 

pectinatae 

costata 

pillcornis 

stroyani 

bogdanowi 

abieticola 

juniperi 

fresai 

cupressi 

tujafilina 



Body 
length 



4-6-6-4 

4-8-5*5 

3.4-4-6 

3-1-4-8 

3-2-4-1 

2-2-4-0 

3'°-4'4 

3'4-4*4 

2-8-37 

3 '3-5-3 

3"4-4 -6 

3'3-5-o 

3-0-4-4 

2-6-4-4 

2-0-3-8 

3-2-4-7 

5-i-7'5 

2-6-3-1 

2-7-4-2 

2-2-3-5 

2-5-4-1 



Diameter 

of 

siphuncular 

cone 

410-950 

420-440 

250-450 

300-650 

360-530 

240-430 

240-370 

300—490 

440-620 

370-730 

450-580 

150—400 

320-600 

170-370 

130-390 

500—670 

500-990 

330-460 

250-600 

300-450 

170-380 



Chitinised Scleroites 

rim of on abdominal 

primary 
rh in aria 



evident 

present 
evident 



Length of antennal segments 



narrow 
present 



evident 

absent 



tergites 






2-5 I" 


IV 


V 


small or absent 650-1230 


290-420 


460-5S0 


820-870 


400-460 


500-520 


small 55o-75o 


250-330 


340-420 


evident 500-75° 


220-340 


280-420 


absent 560-650 


210-290 


260-320 


440-700 


1S0-300 


220-330 


absent or v. small 550-75° 


200-340 


250-37° 


,. .. 610-700 


250-320 


320-360 


absent 520-650 


220-290 


270-320 


large 54°-750 


230-350 


300—420 


small 590-730 


270-310 


290-340 


small to large 300-550 


160—200 


180-230 


v. small to absent 400-660 


170-240 


210-260 


absent 350-5°° 


130-240 


160-280 


360-480 


140—200 


160-250 


"„ 540-850 


240-430 


210-460 


v. small 800-1150 


380-620 


3S0-650 


360-460 


130—210 


190-270 


390-560 


160-280 


200-290 


v. small or absent 35°-55° 


160-220 


180-230 


„ » 3i°-4 10 


140-200 


160-190 



VI 

190-210 + 50-70 
150-180 + 65-90 
150-190 + 25-40 
110-160 + 25-45 
120-160 + 35-50 
110-160 + 35-6° 
120-160 + 35-50 
140-170 + 40-55 
170-220 + 60-90 
150-230 + 40-85 
150-190 + 40-70 
160-200 + 29-50 
160-220 + 14-40 
1 10-190 + 25-55 
120-170 + 25-35 
150-210 + 25-70 
250-320 + 45-75 
180-230 + 40-75 
150-240 + 38-75 
j 40-1 90 + 30-50 
130-180 + 14-30 



Length 


of 


rostral segments 


4 


5 


300-370 


130-150 


330-360 


140-150 


160-190 


90-105 


160-200 


85-110 


190-230 


1 10-135 


140-180 


70-100 


130-170 


80-100 


200-220 


95-110 


260-300 


130-145 


210-270 


iiS-^o 


3i°-330 


120-170 


190-250 


90-100 


190-250 


95-125 


160-250 


95-140 


180-240 


95-120 


230-290 


105-135 


290-370 


130-165 


130-160 


60-95 


140-210 


85-115 


130-170 


75-100 


130-180 


65-100 





Hind tarsus 




Length 
of 


first segment 


length 


hind 






of 
second 


tibia 


dorsal 


ventral 


in 


length 


length 


segment 


mm 


65-100 


170-220 


450-520 


4-3-6-3 


75-85 


170-200 


410-440 


3-5-3-9 


90-120 


180-220 


440-510 


2-6-3-2 


60-90 


130-180 


320-400 


2-2-3-0 


35-55 


100-130 


250-310 


2-5-3-2 


35-°° 


90-125 


220-300 


1-8-2-9 


45-75 


110-160 


260-330 


2-3-2-9 


45-75 


130-155 


300-360 


2-9-3-2 


80-110 


160-190 


350-410 


2-1-2-5 


130-240 


210-330 


340-520 


2-2-3'2 


70-120 


160-200 


320-360 


2-4-3-1 


150-180 


190-270 


380-450 


I-4-2-2 


25-50 


100-120 


300-370 


2-0— 2-6 


25-55 


95-^55 


35°-54° 


1-5-2-2 


35-55 


100-125 


320-340 


1-4-2-1 


50-85 


125-180 


310-340 


2-4-3*5 
4.3-6-7 


30-55 


1 40-200 


420-570 


20-35 


85-110 


320-370 


l-6-i*9 


15-40 


90-110 


280-390 


1-6-2*6 


20-35 


80-110 


280-350 


1-4-2-2 

1-6-2-1 


25-35 


90-110 


240-340 



Table 2 (contd.) 



Length of longest hairs on 



kochiana 

boerneri 

iaricis 

acutirostris 

pini 

pinihabitans 

escherichi 

brauni 

pinea 

schimitscheki 

pectinatae 

costata 

pilicornis 

stroyani 



third hind 

antennal tibia 
segment 



40-65 

55-70 

25-40 

50-80 

70-100 

50-90 

120-160 
55-75 
80-110 

120-170 

120-150 
75-110 

150-210 
80-160 
70-105 



45-75 
45-75 
50-80 
120-150 
1 10-130 

75-I40 
170-210 

75-95 
140-160 
190-270 
150-170 
190-240 
190-290 
180-270 

95-180 



Abdominal 

tergites 
3 8 



35-So 
30-60 
20-40 
70-110 
65-100 
15-75 
120-200 
3°-55 
120-180 
150-240 
70-110 
85-15° 
130-160 
105-160 
55-no 



30-140 
80-100 
75-100 
130-190 
120-160 
90-150 
150-200 
85-120 
130-200 
170-250 
1 10-150 
170-210 
150-210 
125-220 
100-200 



Number of 
hairs on 
antennal 
segments 

II VI 

Base 

11-18 12-20 

21-32 15-22 

8-13 7-9 



Number of hairs on 



5-9 
6-S 
6-9 
6-9 



4-7 
9-13 
8-14 

7-1 1 



7-13 12-16 
7-12 11-14 
3-io 3-9 

7-13 7-8 
6-8 5-7 

11—17 12—20 
8-20 6-13 

9-18 9-17 



sub-apical accessory hairs 
processus on 

terminalis rostral segment 



7-1 1 

5-8 
4(-5) 
(3")4 

(3-)4(-5) 
(3-)4 

6-7 
6-7 
2-5 
(3-)4 
3-4 
3-4 
3-4 
4-5 



140-190 160-240 130-190 150-210 14-24 11-19 4 

220-320 220-380 160-320 250-320 18-25 10-14 (3 - )4 

160—200 250-320 160-240 220-260 7-11 6-12 3(~4) 

190-250 280-360 140-210 190-260 10-14 8-14 (2-)3(-4) 

160-220 230-350 120-200 160-260 7-10 5-6(-7) (2-)3 - 

120-200 140-310 120-210 130-210 9-1 1 g-13 3{-4) 



10-17 
2 4-34 
4-8 
5-8 
6-8 
6-10 
4-5 
8-1 1 
4-7 
4-6 
6-7 
7-8 
4-6 
4-6 



8-15 
8-13 



5-8 
2-5 



Sub- 


Abdominal 


Secondary rhinaria on 


genital 


tergites 


antennal segments 


plate 
























5 


8 


III 


IV 


V 


60-110 


6-14 


12-20 


5-15 


1-3 


1-2 


33-55 


18-40 


43-61 


11-14 


2-5 


2-4 


35-48 


19-24 


16-32 


7-13 


i-4 


1-3 


12-29 


16-30 


14-25 


3 -1 * 


0-4 


(0-)l(-2) 


24-38 


5-6 


10-17 


1-6 


1-4 


I 


20-42 


4-7 


9-19 


5-10 


1-3 


1-2 


17-29 


18-38 


11-22 


1-8 


1-4 


1-2 


39-58 


4-6 


17-22 


4-10 


2-4 


1-3 


30-62 


16-26 


12-16 


g-i 7 


2-4 


i-3 


29-49 


18-39 


17-27 


1-18 


0-5 


0-3 


50-70 


50-58 


35-43 


o-9 


0-1 


1 


40-70 


37-5° 


24-50 


0-9 


0-6 


0— 1 


29-43 


32-68 


26-42 


1-3 


1-2 


0-2 


18-42 


24-49 


21-53 


1-8 


0-3 


1-2 


22-37 


24-49 


16-32 


6-9 


1-3 


1-2 


22-36 


30-55 


24-42 


1-8 


1-3 


1 


32-45 


50-70 


29-42 


7-15 


2-5 


0-3 



23-34 2 5-3° T 9-Z4 4-8 J ~4 0-2 

26-40 60-70 16-27 6-1 1 1-4 1-2 

22-40 60-70 15-25 3-6 1-3 1-3 

2 4-3S 38-60 17-24 3-8 1-3 1-2 



Other characters 



Proximal 2/3 of hind femur pale 



Hind femur pa! 
Hind tibiae 



near base only 
h a pale area near base 



Hind tibiae black 

Hind tibiae dark near apex 

Fore-wing maculate near apex 

Hind tibiae dark near apex 

Hind tibiae dark or with a very small dusky area near 

the base 

Hind tibia with pale area near base 

Hind tibiae black or with a small brown area near the 

base 

Hind tibiae black 

Hind tibiae dark at base and apex 
Hind tibiae dark at apex only 



REVIEW OF CINARA IN BRITAIN 



173 



Table 3 

Discriminants for the apterae viviparae of the British species of Cinara 





Third antenna] 


segment 


Hind 






Rostral 
segment 


Hind 
tarsal 












longest 


tibia: 


Antennal 


Rostral 


4: 


segment 




diameter of 


hair 


longest 


segments 


segments 


hind 


2: 




siphuncular 


borne 


hair 


V : VI 


4 : 5 


tarsal 


rostral 




cone 


on it 


borne 
on it 






segment 
1 


segment 
4 


piceae 


1-0-3-2 


7-5-31 


36-84 


1 -6-2- 1 


2-2-3-0 


1-8-2-5 


i-i-i-3 


fundatrix 


1-8-2-4 


12-23 


57-79 


1-6-1-9 


2-2-2*9 


1-6-2-0 


1-1-1-4 


kochiana 


1-6-2-2 


8-5-15 


38-78 


1-7-2-2 


2-4-3-0 


1-8-2-2 


I-0-I-2 


boerneri 


1-8-3-5 


7-8-25 


32-56 


1-4-1-9 


1-7-2-3 


0-9-1-3 


1-8-2-6 


laricis 


0-8-2-1 


6-8-13 


17-34 


1-4-2-5 


1-2-2-2 


1-0-1-7 


1-4-2-1 


acutirostris 


o-8-i-6 


6-6-9-3 


24-33 


I-3-I-8 


1-8-2-3 


I-6-2-I 


1-0-1-4 


fundatrix 


I-2-I-6 


5-0-6-0 


16-19 


1-1-1-4 


1-7-1-9 


1-6-2-0 


1-1-1-3 


pini 


0-7-2-5 


7-7-15 


23-42 


1-1-1-9 


1-7-2-3 


1-3-1-9 


1-2-1-7 


fundatrix 


0-9-1-6 


6-6-1 1 


22-39 


I-2-I-8 


1-7-2-2 


I-2-I-6 


I-5-I-8 


pinihabitans 


I-2-I-6 


4-4-6-0 


14-16 


1-4-1-8 


1-7-1-9 


1-0-1-3 


1-7-2-3 


escherichi 


1-1-1-7 


9-5-15 


30-46 


1-4-1-8 


2-0-2-4 


1-1-1-7 


1 -2-1-5 


fundatrix 


0-9-1-5 


9-5-1 1 


31-34 


i-i-i-3 


2-0-2-1 


I-3-I-5 


I-6-I-8 


brauni 


0-8-1-2 


4-5-9-5 


16-25 


0-9-1-3 


1-8-2-5 


1-5-2-0 


1-0-1-5 


pinea 


0-8-2-1 


3-2-6-5 


12-20 


1-1-1-7 


1-6-2-2 


0-8-1-2 


1-4-2-0 


schimitscheki 


1-0-1-5 


5-9-12 


15-27 


I-3-I-5 


2-1-2-4 


i -4-1 -8 


0-9-1-2 


pectinatae 


1-4-2-2 


3-4-" 


7-18 


0-7-1-0 


2-4-2-9 


I-0-I-2 


1-5-1-9 


costata 


0-7-1-2 


2-0-2-8 


6-9 


0-8-1-2 


1-8-2-4 


1-7-2-5 


I -2-1-6 


pilicornis 


1-2-2-2 


2-3-4-5 


7-12 


1-0-1-4 


I-8-2-I 


1-4-2-3 


1-5-2-4 


stroyani 


0-6-2-3 


4-2-7-5 


17-29 


1-1-1-4 


2-1-2-5 


1-5-2-3 


1-2-1-9 


fundatrix 


1-2-2-3 


4-7-6-7 


14-21 


0-9-1-4 


2-2-2-4 


1-7-2-1 


1-4-1-6 


bogdanowi 


O-6-I-g 


3-2-6-2 


14-27 


1-0-1-7 


1-9-2-6 


1-6-2-3 


0-9-1-3 


fundatrix 


0-7-0-9 


3-6-4-7 


11-18 


1-0-1-5 


2-3-2-6 


1-8-2-3 


0-9-1-3 


abieticola 


0-9-1-9 


2-6-4-4 


12-29 


I-2-I-8 


2-1-2-6 


1-8-2-4 


1-2-1-5 


juniperi 


0-5-1-1 


J-5-2-3 


4-8-3 


0-7-1-1 


I-6-2-I 


1-4-2-1 


1-8-2-4 


fresai 


0-7-1-7 


1-7-2-4 


6-9 


0-7-1-1 


1-6-2-4 


1-8-2-2 


1-3-1-7 


cupressi 


1-1-1-7 


1-6-2-5 


5-8 


0-7-1-1 


1-5-2-1 


1-4-1-9 


1-5-2-0 


tujafilina 


o-8-i-5 


1-6-2-5 


5-9 


o-8-i-i 


1-8-2-2 


I-6-2-I 


1-4-1-7 



174 



V. F. EASTOP 



Table 4 

Discriminants for the alatae viviparae of the British species of Cinara 





Third antennal 


segment 


Hind 






Rostral 
segment 


Hind 
tarsal 
















longest 


tibia : 


Antennal 


Rostral 


4: 


segment 


Hind 




diameter of 


hair 


longest 


segments 


segments 


hind 


2: 


tarsal 




siphuncular 


borne 


hair 


V : VI 


4 : 5 


tarsal 


rostral 


segments 




cone 


on it 


borne 
on it 






segment 
1 


segment 
4 


2 : 1 


piceae 


1*1-2*1 


17-26 


65-120 


1-8-2-3 


2-2-2-8 


1-5-1-9 


1-3-1-6 


2-2-2-7 


kochiana 


1*9-2-0 


11-15 


50-75 


1-9-2-3 


2-3-2-4 


1-7-2-0 


1-2-1-3 


2-1-2-4 


boerneri 


1-3-2-6 


17-25 


38-60 


1-7-2-1 


I-7-2-I 


o-8-i-o 


2-4-2-8 


2*3-2*7 


laricis 


1-0-1-9 


7-12 


16-23 


1-7-2-7 


I-6-2-I 


1-0-1-3 


1-8-2-4 


2*1-2*7 


acutirostris 


i-i-i-6 


6-8-5 


20-29 


1-4-1-9 


I-6-I-8 


1-7-2-1 


1-1-1-4 


2*3-2-6 


pini 


1-3-2-3 


5-5-9 


14-25 


1-2-1-7 


1-6-2-2 


I-2-I-6 


1-5-1-8 


2-2-2-7 


pinihabitans 


1-8-2-6 


3-5-6 


12-16 


1-5-2-0 


1-5-1-9 


1-0-1-3 


1-7-2-2 


2-0-2-4 


escherichi 


1 -3-2- 1 


8-5-12 


32-38 


1-5-1-9 


2-0-2-2 


1-3-1-6 


1-4-1-7 


2-1-2-6 


brauni 


1-0-1-3 


6-5-8 


14-16 


I-0-I-2 


1-9-2-3 


I-5-I-8 


1-2-1-5 


2-0-2-3 


pinea 


0-9-1-7 


3-5-6 


9-5-16 


I-I-I-7 


I -6-2- 1 


0-7-1-1 


1-5-2-1 


I-4-2-0 


schimitscheki 


1-2-1-5 


4-6 


15-19 


1-2-1-5 


1-9-2-6 


1-6-2-0 


I-0-I-2 


1-8-2-0 


pectinatae 


I-I-2-O 


3-6-5 


6-12 


o-8-i-i 


2-1-2-8 


o-8-i-i 


I-7-2-I 


I-7-2-I 


costata 


0-8-1-5 


2-4-3-3 


7*5-i° 


0-9-1-3 


I-7-2-2 


1-7-2-3 


1-3-1-7 


2-8-3-5 


pilicornis 


1-2-2-2 


3-5 


6-5-1 1 


0-9-1-4 


I-6-2-I 


1-4-2-0 


1-8-2-6 


3-2-4-4 


stroyani 


1-2-2-2 


3-7-6-1 


9-5-18 


i-c-i-3 


1-9-2-2 


1-6-2-2 


1-6-1*9 


3-1-3-5 


bogdanowi 


O-g-I-6 


2-9-5-1 


11-23 


1-5-1-9 


1-9-2-6 


1-5-1-9 


I-I-I-5 


2-2-2-7 


abieticola 


1-1-1-9 


2-9-4-5 


14-24 


I-3-I-8 


2-0-2-5 


1-6-2-5 


1-4-1-6 


2-6-3-5 


juniperi 


O-g-I-2 


1-9-2-6 


5-7-6-8 


0-7-1-1 


1-5-2-0 


1-4-1-6 


2-0-2*8 


3-3-4-° 


fresai 


o-8-i-6 


1-8-2-4 


5-8-7-7 


o-8-i-i 


1-6-2-2 


I-6-2-I 


1*6-2*1 


2*9-3-5 


cupressi 


1-0-1-5 


2-1-2-6 


5-6-6-8 


0-9-1-2 


1-5-1-9 


1-2-1-7 


2-0— 2*4 


2-8-3*5 


tujafilina 


1-0-1-9 


2-0-2-9 


6-3-8-6 


0-9-1-1 


1-5-2-2 


I-2-I-8 


1-6-2-2 


2*6-3*3 



REVIEW OF CINARA IN BRITAIN 



175 



Table 5 
Biometric and other data for the sexuales of the British species of Cinara 

u "a Males 

0) CD 

Available -g .5 Secondary rhinaria on antennal Pseudosensoria 

subgeneric Condition 3 | segments on hind tibiae o 

name ^ £ III IV V 



piceae 
kochiana 

boerneri 
laricis 

acutirostris 

pint 

pini subsp. 

montanicola 
pinihabitans 
escherichi 
brauni 
pinea 

schimitscheki 
pectinatae 
costata 

pilicornis 

stroyani 

bogdanowi 

abieticola 

juniperi 

Jresai 

cupressi 

tujafilina 



Mecinaria 
Laricaria 

Cinarellia 
Cinaria 

Cinara 



Subcinara 
Cinarella 

Buchneria 
Lachniella 

Cinaropsis 



unknown 
alate 



apterous 
alate 



unknown 
alate 



Pityaria 

Dinolachnus 

Cupressobium unknown 

alate 
unknown 



IV V VI 

I 90-98 18-20 12-15 



oviparae 



— absent or inconspici 
o numerous, 

inconspicuous 
15 82-1 1 1 19-39 13-20 9-14 numerous 
8 60-110 18-31 11-19 6-11 numerous, 

inconspicuous 
2_ 5 7~8 absent 



apterous 3 

alate 1 

? apterous o 
unknown o 
alate 
unknown 
alate 
alate 



0-2 
33-37 



5-8 
7-1 1 



3-27 3-8 
91-116 20-30 



2-5 

2-5 
7-8 



45-63 n-24 7- IQ 



o inconspicuous 

o absent or inconspici; 

o conspicuous 

— unknown 



o numerous, conspicuc 
— unknown 



56-68 
38-60 

45-58 



14-27 12-14 4 _ 5 conspicuous 
13-25 6-12 

13-18 3-4 



46-82 12-26 4-15 o 
102-130 35-39 14-1 8 ° 



o conspicuous, tibia 

swollen 
o numerous, conspicu< 

indistinct 

numerous, conspicuc 



22-39 



i-12 



6-7 



unknown 

numerous, conspicuc 
unknown 



176 V. F. EASTOP 

REFERENCES 

Aizenberg, E. E. 1956. New contribution to aphid taxonomy (Homoptera) . Trudy vses. 

ent. Obshch. 45 : 128-166. [In Russian.] 
Altum, B. 1882. Forstzoologie. III. Insecten. 2. Aufl. 2. Abth. Berlin, 382 pp. [aphids 

PP- 350-358]- 
Archibald, K. D. 1958. Forest Aphididae of Nova Scotia. Proc. Nova Scotian Inst. Sci. 

24 (2) : 1-254. 
Arnhart, L. 1927. Osterreichischer Larchenhonigtau, Larchenmanna und Larchenhonig. 

Z. angew. Ent. 12 : 457-472. 

1930. Die Latschen-Honigtau liefernde Lachnus Nenbergi n. spec. Z. angew. Ent. 16 : 

392-398. 

Baker, A. C. 1919a. On the use of the names Lachnus and Lachniella. Can. Ent. 5 : 211- 

212. 
19196. Wilsonia — a correction. Can. Ent. 5 : 253. 

1920. Generic Classification of the Hemipterous Family Aphididae. Bull. U.S. Dep. 

Agric. 826 : 109 pp. 

Blanchard, E. E. 1939. Estudio sistematico de los Afidoideos argentinos. Physis 17 : 

857-1003. 
Bodenheimer, F. S. & Swirski, E. 1957- The Aphidoidea of the Middle East. Weizmann 

Science Press, Jerusalem, 378 pp. 
Borner, C. 1932. Author responsible for the systematic part of Borner & Schilder, 1932, see 

footnote 7 on page 551. 
1939. Neue Gattungen und Arten der mitteleuropaischen Aphidenfauna. Arb. physiol. 

angew. Ent. Berl. 6 : 75-83. 

1940. Neue Blattlduse aus Mitteleuropa. Privately published, Naumburg (Saale), 4 pp. 

1949- Kleine Beitrage zur Monographic der europaischen Blattlause. Beitr. tax. Zoo!. 

1 : 44-62. 

1950. Neue europdische Blattlausavten. Privately published, Naumburg (Saale), 19 pp. 

1952 & 1953. Europae centralis Aphides. Mitt, thiiring. bot. Ges. 4 (3) 1 & 2 : 1-259, 

260-484 (1952), 485-488 (1953)- 

Borner, C. & Franz, H. 1956. Die Blattlause des Nordostalpengebietes und seines Vor- 
landes. Ost. zool. Z. 6 : 297-411. 

Borner, C. & Heinze, K. 1957. Aphidina — Aphidoidea in Sorauer, P., Handbuch der 
Pflanzenkrankheiten 5 (2) Homoptera II. Berlin & Hamburg, 577 pp., 5th ed. [aphids 
pp. 1-402]. 

Borner, C. & Schilder, F. A. 1932. Aphidina in Sorauer, P., Handbuch der Pflanzenkrank- 
heiten 5 (2). Berlin, 1032 pp., 4th ed. [Aphidina pp. 551-715.] 

Boudreaux, H. B. 1948. New species of Louisiana Aphididae, and notes on Sanbornia 
juniperi Pergande. Fla Ent. 31 : 96-105. 

Bradley, G. A. 1951. A Field Key to the Species of Cinara (Homoptera: Aphididae) of 
Canada East of the Rockies. Can. Ent. 83 : 333-335. 

1956. Three new Species of the Genus Cinara Curtis (Homoptera: Aphididae) from 

Larix spp. Can. Ent. 88 : 492-495. 

1959. Feeding Sites of Aphids of the Genus Cinara Curtis (Homoptera: Aphididae) in 

North Western Ontario. Can. Ent. 91 : 670-671. 

1963. Two New Species of Cinara Curtis (Homoptera: Aphididae) from Juniperus 

horizontalis Moench. Can. Ent. 95 : 287-291. 

• 1965. A New Species of Cupressobium (Homoptera: Lachnidae) from Hawaii. Can. 

Ent. 97 : 668-670. 
Bradley, G. A. & Hinks, J. D. 1968. Ants, Aphids and Jack Pine in Manitoba. Can. 

Ent. 100 : 40-50. 
Braun, R. 1938. Die Honigtaufrage und die honigtauliefernden Kienlause (Cinarini C.B.). 

Z. angew. Ent. 24 : 462-510. 



REVIEW OF CINARA IN BRITAIN i 77 

Bray, D. F. 1953- Life History and Control of Cinara winokae. J . econ. Ent. 46 : 103-107 

Buckton, G. B. 1876-1883. Monograph of the British Aphides. Ray Society, London. 
Vol. 1, 193 pp. + plates A-C & 1-38 (1876); vol. 2, 176 pp. + plates 39-86 (1879); vol. 3, 
142 pp. + plates 87-114 (1881); vol. 4, 228 pp. + plates D-I & 115-134 (1883). 

Burmeister, H. 1835. Handbuch der Entomologie. 2 (i) Rhynchota. Berlin, 400 pp. [aphids 
PP- 85-95]. 

Burnham, J. C. 1938. A contribution to the List of Aphididae of the Maritime Provinces of 
Canada. Can. Ent. 70 : 180-188. 

Canakcioglu, H. 1966. A study of the forest Aphidoidea of Turkey. Istanb. Univ. Orman 
Fak. Derg. Ser. A 16 (2) : 1 31-190. [In Turkish with English summary]. 

Cholodkovsky, N. A. 1892. Wissenschaftliche Mittheilungen. 1. Zur Kenntnis der 
Coniferen-Lause. Zool. Anz. 15 : 74-78. 

1896a. Zur Kenntnis der auf Fichte (Picea excelsa Lk.) lebenden Lachnus- Arten. Zool. 

Anz. 19 : 145-150, 200 (Berichtigungen) . 

18966. Aphidologische Mittheilungen. I. Uber die auf Nadelholzern lebenden Lachnus- 
Arten. Zool. Anz. 19 : 508-509. 

1898. Beitrage zu einer Monographic der Coniferen-Lause. Trudy russk. ent. Obshch. 

31 : 603-674 + plates xi-xiii. 

1899. Aphidologische Mittheilungen. 7. Uber einige neue oder wenig bekannte 

Lachnus- Arten. Zool. Anz. 22 : 468-473 + plate. 

1902. Second catalogue of the collection of aphids (Aphidae) in the zoological cabinet 

of the St. Petersburg Forestry Institute 8 (2) : 1-11. [Reprint, in Russian.] 

Curtis, J. 1835. British Entomology. 12, parts 576 & 577, each 2 pp. + plate. London. 
Dale, H. C. 1969. The distal apical structure of aphid siphunculi (Hemiptera, Aphididae). 

Entomologist' s Gaz. 20 : 270 + plates 2 & 3. 
Daoud, A. A. K. & El-Haidari, H. 1968. Recorded Aphids of Iraq. Iraq nat. Hist. Mus. 

Pubis 24 : 37 pp. + map. 
1969. Some new records of aphids from Iraq. Bull. Iraq nat. Hist. Mus. 4 (1) : 

10-14. 
Davidson, W. M. 1914. Plant-louse notes from California. /. econ. Ent. 7 : 127-136. 
de Geer, C. 1773. Memoires pour servir a I'histoire des Insectes 3 : viii + 697 pp. [Aphis. 

pp. 19-129 & plates 2-7) Stockholm. 

in Goeze, J. A. E. 1780. Abhandlungen zur Geschichte der Insekten. Niirnberg, 

454 pp. + 18 pp. of index. 

1783. Genera et Species Insectorum. Leipzig, 220 pp. (aphids pp. 77-78). 

Doncaster, J. P. 1961. Francis Walker's Aphids. British Museum (Natural History), 

London, 165 pp. + 8 plates. 

Dufour, L. 1833. Recherches anatomiques et physiologiques sur les Hemipteres, accom- 
pagnees de considerations relatives a I'histoire naturelle et a la classification de ces in- 
sectes. Mim. pres. div. Sav. Acad. Sci. Inst. Fr. 4 : 131-461 + 19 plates. 

Dzhibladze, A. A. 1958. Contribution to the study of the aphid fauna of Eastern Kakhetia. 
Trudy Inst. Zool., Tbilisi 16 : 291-321 [in Russian]. 

i960. Notes on the Aphidoidea of the western Caucasian mountain ridge. Trudy Inst. 

Zool., Tbilisi 17 : 19-30 [in Russian]. 

Eastop, V. F. 1958. A Study of the Aphididae (Homoptera) of East Africa. H.M.S.O. 
London, 126 pp. 

1 961. A Study of the Aphididae [Homoptera) of West Africa. British Museum (Natural 

History), London, 93 pp. 

1966. A taxonomic study of Australian Aphidoidea (Homoptera). Austr. J. Zool. 

14 : 399-592. 

Essig, E. O. 191 1. Aphididae of Southern California. VII. Pomona Coll. J. Ent. 3 (3) : 

523-557- 
Fabricius, J. C. 1781. Species Insectorum. 2. Hamburg & Kiel. 517 pp. [aphids pp. 
384-390]. 



i 7 8 V. F. EASTOP 

Fabricius, J. C. 1794- Entomologia systematica. 4. Copenhagen. 477 pp. + 175 pp. of 
index [aphids pp. 294-302]. 

1803. Systema Rhyngotorum. Brunswick. 315 pp. + 23 pp. of index [aphids pp. 

294-303]- 
Fitch, A. 1859. Third, Fourth and Fifth reports on the Noxious, Beneficial and other Insects 

of the State of New York. Albany. Third Report. Insects infesting fruit trees, 172 pp. 

Fourth Report. Insects infesting Evergreen Forest Trees, 67 pp. Fifth Report. Insects 

infesting Deciduous Forest Trees, 74 pp. + Index to all three reports, 11 pp. 
Fossel, A. 1958. Die Tannentracht. Bienenvater, Wien 79 : 163-169, 197-202. 

i960. Die Fichtentracht. Bienenvater, Wien 81 : 204-228. 

Froggatt, W. W. 1923. Forest Insects of Australia. Forestry Commission of New South 
Wales, Sydney, 171 pp. 

1927. Forest Insects and Timber Borers. Sydney, 107 pp. 

Gillette, C. P. & Palmer, M. A. 1924. New Colorado Lachnini. Ann. ent. Soc. Am. 
17 : 1-44 + 13 plates. 

1930. Three new aphids from Colorado. Ann. ent. Soc. Am. 23 : 543-551. 

1931. The Aphididae of Colorado. Part 1. Ann. ent. Soc. Am. 24 : 827-934. 

Gomez Menor Ortega, J. 1934. Insectos que atacan al Pino en la Republica. Rev. Agric. 

Com. S. Domingo 25 : 1372-1373. 

1962. Lachnidae de Espana (Horn. Sternorrhyncha) . Eos, Madr. 38 : 347-413. 

Gontarski, H. 1941. Beitrag zur Honigtaufrage. Z. angew. Ent. 27 : 321-332 (1940). 
Goot, P. van der. 1915. Beitrdge zur Kenntnis der Holldndischen Blattlduse. Haarlem 

& Berlin, viii + 1 p. errata + 600 pp. + 8 plates. 

1918. Zur Kenntnis der Blattlause Java's. Contr. Faune Indes Neerl. 3 : 1-301. 

Grechkin, V. P. 1962. Some pests of young coniferous trees in Eastern Siberia. Zool. 

Zh. 46 : 706-716 [in Russian with English summary], 
Guercio, G. del. 1900 & 1902. Prospetto dell'Afidofauna Italica. Osservazioni preliminari. 

Nuove Relaz. R. Staz. Ent. agr. (s. 1) no. 2 : 1-172 + plate (1900); pp. 173-236 (Cher- 

mesidae) (1902). 
1909. Contribuzione alia conoscenza dei Lacnidi Italiani. Redia 5 : 173-359 + plates 9- 

20. 
Gunkel, W. 1963a. Cupressobium juniperinum M.oxd\v. (Homoptera-Lachnidae), ein Schad- 

ling an Thuja occidentalis L. Beitrag zur seiner Morphologie und Biologie. Z. angew. 

Zool. 50 : 1-48. 

1 9636. Natiirliche Feinde und Massenwechsel der Lebensbaumblattlaus Cupressobium 

juniperinum Mordw. (Homoptera-Lachnidae). Z. angew. Zool. 50 : 329-341. 

Hall, W. J. 1926. Notes on the Aphididae of Egypt. Bull. Minist. Agric. Egypt tech. scient. 

Serv. 68 : viii + 62 pp. 
Hartig, T. 1839. Jahresberichte iiber die Fortschritte der Forstwissenschaft und forstlichen 

Naturkunde im Jahre 1836 und 1837. Berlin, vol. 1, part 4, Entomologische Notizen, 

pp. 640-646. 

1841. Versuch einer Eintheilung der Pflanzenlause (Phytophthires Burm.) nach der 

Flugelbildung. Z. Ent. (Germar) 3 : 359-376. 

Heie, O. 1964. Aphids collected in Iceland in August, 1961. (Homopt., Aphididae.) 

Ent. Meddr 32 : 220-235. 
■ 1967. Studies on Fossil Aphids (Homoptera: Aphidoidea). Spolia Zool. Mus. haun. 

26 : 273 + 1 p. of addenda. 

1969. The Baltic Amber Aphidoidea of the Geologisches Staatsinstitut of Hamburg. 

Mitt. geol. Stlnst. Hamb. 38 : 143-151. 

Heikinheimo, O. 1963. Fur die Finnische Fauna neue Blattlause (Horn., Aphidoidea) II. 

Suomen hydnt. Aikak. 29 : 184-190. 

1968. The Aphid Fauna of Spitsbergen. Suomen hydnt. Aikak. 34 : 82-93. 

Heinze, K. 1962. Pflanzenschadliche Blattlausarten der Familien Lachnidae, Adelgidae 

und Phylloxeridae, eine systematisch-faunistische Studie. Dt. ent. Z. (n.s.) 9 : 143-227. 



REVIEW OF CINARA IN BRITAIN 179 

Higuchi, H. & Miyazaki, M. 1969. A tentative catalogue of host plants of Aphidoidea in 

Japan. Insecta mats. Suppl. 5 : 1-66. 
Hille Ris Lambers, D. 1931. A list of the Aphididae of Venezia Tridentina, Part I. 

Memorie Mus. Stor. nat. Venezia trident. 1 (1) : 3-5. 

1935- Katalog der Aphiden der Venezia Tridentina. III. Memorie Mus. Stor. nat. 

Venezia trident. 3 : 59—64. 

1948. On Palestine Aphids, with Descriptions of new subgenera and new species (Hom- 

optera, Aphididae). Trans. R. ent. Soc. Lond. 99 : 269-289. 

1952. The aphid fauna of Greenland. Meddr Gronland 136 : 1-33. 

1956. On Aphids from the Netherlands with descriptions of new species. (Aphididae, 

Homoptera.) Tijdschr. Ent. 98 : 229-249. 

1966. Some synonyms in Aphididae (Homoptera). Ent. Ber., Amst. 26 : 124-126. 

Hottes, F. C. 1930. The name Cinara versus the name Lachnus. Proc. biol. Soc. Wash. 

43 : 185-188. 
1934- Aphid descriptions and notes. Proc. biol. Soc. Wash. 47 : 1-8. 

1949- Some obscure aphid species. Proc. biol. Soc. Wash. 62 : 159-160 [Vol. 63 on re- 
print]. 

1953- Notes on some species of Cinara, with descriptions of two new species from Pinon 

Pine. Proc. biol. Soc. Wash. 66 : 153-158. 

1954- Descriptions and notes on some species of Cinara (Aphidae). Proc. biol. Soc. 

Wash. 67 : 251-261. 

*955 a - Cinara descriptions. (Aphidae.) Proc. biol. Soc. Wash. 68 : 67-77. 

19556. Three new subspecies and figures of five previously unfigured species of Cinara 

(Aphidae). Proc. biol. Soc. Wash. 68 : 101-104. 

1958. A new species of Cinara (Aphidae) from Sitka Spruce. Proc. biol. Soc. Wash. 

71 : 61-62. 

1960a. Notes on and a key to the species of Cinara (Family Aphidae) living on Pinus 

edulis. Proc. biol. Soc. Wash. 73 : 199-214. 

iq6o£>. Notes on and key to species of Cinara (Family Aphidae) which have Abies sp. as 

host. Proc. biol. Soc. Wash. 73 : 221-234. 

1961a. Notes on and a key to the species of the genus Cinara (Aphidae) which have 

Tsuga and Pseudotsuga for host. Proc. biol. Soc. Wash. 74 : 111-118. 

19616. A review and key of North American Cinara (Homoptera: Aphididae) occurring 

on Picea. Gt Basin Nat. 21 : 35-50. 

1964. Three new species of Cinara, together with a preliminary list of the species of this 

genus known from Alaska. (Aphididae, Homoptera.) Ent. Ber., Amst. 24 : 50-54. 

& Bradley, G. A. 1953. Two new species of Cinara (Homoptera: Aphididae) 

from Ontario. Proc. biol. Soc. Wash. 66 : 85-88. 

& Essig, E. O. 1953. Descriptions of a new species of Cinara from Western United 

States. Proc. biol. Soc. Wash. 66 : 159-172. 

1954. A new species of Cinara with notes on some Western species of Aphididae. 

Proc. biol. Soc. Wash. 67 : 93-98. 
& Frison, T. H. 1931. The Plant Lice, or Aphididae, of Illinois. Bull. III. nat. Hist. 

Sur. 19 : 121-447. 
Ilharco, F. A. 1968a. Algumas correccoes e adicoes a lista de Afideos de Portugal Continental. 

I Parte (Homoptera- Aphidoidea) . Agronomia lusit. 29 : 11 7-1 39. 

19686. Algumas correccoes e adicoes a lista de Afideos de Portugal Continental. Ill 

parte. (Homoptera-Aphidoidea.) Agronomia lusit. 29 : 247-265. 

1970. Notes on the aphid fauna of Mozambique. Part 1. (Homoptera, Aphidoidea.) 

Rev. Ciencias Biologicas 2 : 1-9. 

Inouye, M. 1937. O ne new an d two unrecorded species of Aphididae from Japan. Insecta 
matsum. 11 : 100-105. 

1939. On eight conifer aphids occurring in Hokkaido. Insecta matsum. 13 : 132-142. 



i8o V. F. EASTOP 

1956. Beitrage zur Kenntnis der Koniferen-lause, vorkommend im nordlichen Teil 

Japans. Spec. Rep. Hokkaido Govt Forest Res. Stn 5 : 204-238. 

— ■ — 1962. Studies on the Scientific Names of Larch-Infesting Aphids. Bull. Govt Forest 
Exp. Stn Meguro 139 : 135-161 + 5 plates. 

1970. Revision of the Conifer Aphid Fauna of Japan (Homoptera: Lachnidae). Bull. 

Govt Forest Exp. Stn Meguro 228 : 57-102 + 18 plates. 

Jackson, D. J. 1919. Further notes on aphides collected principally in the Scottish High- 
lands. Scott. Nat. 93, 94 : 157-165. 

Kaltenbach, J. H. 1843. Monographie der Familien der Pflanzenlduse. (Phytophthires.) 
Aachen, 222 pp. + ip. corrections -f 1 plate. 

1846. Fiinf neue Species aus der Familie der Pflanzenlduse . Stettin, ent. Ztgl : 169—175. 

Kloft, W. i960. Die Trophobiose zwischen Waldameisen und Pflanzenlausern mit Unter- 
suchungen fiber die Wechselwirkungen zwischen Pflanzenlausen und Pflanzengeweben. 
Entomophaga 5 : 43-54. 

Kunkel, H. & Ehrhardt, P. i960. Beitrag zur Lachnidenfauna Mitteleuropas. 

Beitr. Ent. 10 : 161-168. 

Knechtel, W. K. & Manolache, C. I. 1943. Observations on the systematics of aphids 

new to Rumania (second contribution). Anal. Inst. Cere, agron. 13 : 217-267 [1941, in 

Rumanian with German summary]. 
Koch, C. L. 1854-1857. Die Pflanzenlduse Aphiden getreu nach dem Leben abgebildet und 

beschrieben. Nfirnberg, pp. i-vi, 1-134 (1854); 135-236 (1855); 237-274 (1856); vii-viii, 

275-335 ( l8 57) + 54 plates. 
Kring, J. B. 1955- Some Aphid and Host Plant Records from Missouri. /. Kans. ent. 

Soc. 28 : 64-66. 
Kurir, A. 1964. Erstmaliges Massenauftreten der Koniferenlaus Cinaropsis pilicornis 

Hartig in Osterreich. Zentbl. ges. Forstw. 81 : 139-157. 
Leonhardt, H. 1940. Beitrage zur Kenntnis der Lachniden, der wichtigsten Tannen- 

honigtauerzeuger. Z. angew. Ent. 27 : 208—272. 
Lepiney, J. de & Mimeur, J.-M. 1932. Notes d'Entomologie agricole et forestiere du 

Maroc. M6m. Soc. Sci. nat. Phys. Maroc 31 : 1-195. 
Macnay, C. G. 1953. Outbreaks and new Records. PI. Prot. Bull. F.A.O. 2 : 43-45. 
Mamontova-Solucha, V. O. 1963. New data on the aphid fauna of Ukraine (Homoptera, 

Aphidoidea). Trudy Inst. Zool., Kyyiv 19 : 11-40 [in Ukrainian with Russian summary]. 
1964. Aphids of Ukrainian woodlands. Trudy Inst. Zool., Kyyiv 20 : 52-72 [in Ukrainian 

with Russian summary]. 
Maslen, N. R. 1969. Cinara brauni Borner (Homoptera: Aphididae). A New British 

Record. Entomologist 102 : 228. 
Mimeur, J.-M. 1934. Aphididae du Maroc (Troisieme Note). Mem. Soc. Sci. nat. Phys. 

Maroc 40 : 1— 71. 

1936a. Aphididae (Hem.) d'Espagne. Boln Soc. ent. Esp. 19 : 33-40 [known in reprint 

form, pp. 1-8, only]. 

19366. Aphididae (Hem.) du Maroc (Huitieme note). Bull. Soc. Sci. nat. Maroc. 16 : 

252-255- 
Mordwilko, A. 1895a. Zur Biologie und Systematik der Baumlause (Lachninae Pass. 

partim) des Weichselgebietes. Zool. Anz. 18 : 73-85, 93-104. 
18956. On the fauna and anatomy of the fam. Aphididae of the Vistula basin. Rab. 

Lab. zool. Kab. imp. varsh. Univ. 3 : 1-274 + i—viii + i— iv + 2 plates [in Russian]. 

1929. Food plant catalogue of the Aphididae of the U.S.S.R. Trudy prikl. Ent. 14 : 1- 

100 [in Russian]. 

— — 1933- In Philipjev, J. N. & Ogloblin, D. A. Keys to Insects. Moscow & Leningrad 

(Aphidoidea pp. 147-175) [in Russian]. 
— — 1948. In Tarbinski & Plavilshchikov. Keys to the insects of the European Part of 

the U.S.S.R. Aphidoidea pp. 187-226. Moscow & Leningrad [in Russian]. 



REVIEW OF CINARA IN BRITAIN 181 

Mound, L. A. 1969. A species of aphid newly recorded in Britain. Entomologist's mon. 

Mag. 105 : 62. 
Muller, F. P. & Scholl, S. E. 1958. Some notes on the aphid fauna of South Africa. J. 

ent. Soc. sth. Afr. 21 : 382-412. 
Muller, H. i960. Der Honigtau als Nahrung der hiigelbauenden Waldameisen. Ento- 

mophaga 5 : 55-75- 
Narzikulov, M. N. 1963. Aphids (Homoptera, Aphididae) of Tadzhikistan and adjacent 

republics of central Asia. Fauna ofTadzhik SSR 9 (1) : 1-272 [in Russian]. 
Nevsky, V. P. 1929. The plant lice of central Asia. Trudy uzbekist. opyt. Sta. Zashch. 

Rast. 16 : 1-425 [In Russian]. 
Nordlinger, A. 1880. Lebensweise von Forstkerfen oder Nachtrdge zu Ratzeburg's Forstinsekten 

2 Aufl. Stuttgart. 
Nordlinger, H. 1864. Waldhonigthau. Krit. Bldtt. Forst.-u. Jagdwiss. 46 (2) : 128-137. 
Nuorteva, P. 1957. Cinara piceae (Panz.) (Horn., Aphididae) found in Finland. Suomen 

hydnt. Aikak. 23 : 35-36. 
— ■ — 1958. On the occurrence of proteases and amylases in the salivary glands of Cinara piceae 

(Panz.) (Horn. Aphididae). Suomen hydnt. Aikak. 24 : 89. 
Oestlund, O. W. 1922. A Synoptical key to the Aphididae of Minnesota. Rep. St. Ent. 

Minn. 19 : 114-151. 

1942. Systema Aphididae. A guide to the phytogeny of the Aphids or plant lice. Part 1. 

The Lachnea. Rock I., Illinois, v + 88 pp. 

Okamoto, H. & Takahashi, R. 1927. Some Aphididae from Corea. Insecta matsum. 

1 : 130-148 + plate 4. 

Orr, M. Y. 1932. An aphid (Neochmosis vanduzei Swain) new to the Forth Area. Scott. 

Nat. 194 : 53. 
Ossiannilsson, F. 1959. Contributions to the knowledge of Swedish Aphids. II. List of 

Species with Find Records and Ecological Notes. K. LantbrHogsk. Annlr 25 : 375-527. 
Paik, W. H. 1965. Aphids of Korea. Seoul National University, 160 pp. 
Palmer, M. A. 1952. Aphids of the Rocky Mountain Region. Thomas Say Foundation, 

Denver, 452 pp. 
Panzer, G. W. F. 1801. Fauna Insectorum Germaniae Inilia. Part 78, no. 22, Nurnberg, 

2 pp. + plate. 

Pasek, V. 1952. The Czechoslovakian Lachnids (Homopt., Aphidoidea). A faunistic 
review. Biol. Sb., Bratisl. 7 : 91-99 [in Czech with German summary]. 

J 953 a - Contribution to the classification of central European Lachnids (Homoptera, 

Aphidoidea). VSst. csl. Spot. zool. 17 : 149-177 [in Czech with German summary]. 

I953&- Some new Lachnids (Homopt., Aphidoidea) from Czechoslovakia. I. Zool. 

ent. Listy 2 : 28-34. 

J 953 c - Cinara doncasteri sp. n. and two more interesting Lachnidae. Rod. 6sl. Spol. ent. 

50 : 222-231 [in Czech with Russian and German summaries]. 

1954- Aphids attacking Coniferous Trees in Czechoslovakian Forests. Slovak Academy 

of Science, Bratislava, 319 + 2 PP- [in Czech]. 

Patch, E. M. 1912. Aphid Pests of Maine. Bull. Me agric. Exp. Stn 203 : 159-178. 
Pettersson, J. 1970. An Aphid Sex Attractant. I. Ent. Scand. 1 : 63-73. 
Pintera, A. 1959. Faunistic contribution to the knowledge of Bulgarian Aphids (Horn., 
Aphid.). Cas. 6sl. Spol. ent. 56 : 69-80 + 2 plates. 

1966. Revision of the genus Cinara Curt. (Aphidoidea, Lachnidae) in Middle Europe. 

Acta ent. bohemoslovaca 63 : 281-321. 

Rihar, J. 1963. Plant Lice and scales (Aphidoidea and Coccidea) on Forest- and Fruit-trees 
as producers of Honeydew in Slovenia. Zast. Bilja 73 : 255-271 [in Serbian with English 
summary] . 

Rupais, A. A. 1961. Dendrophilous aphides in parks and public gardens of the Latvian SSR. 
Latvian Academy of Sciences, Riga, 252 pp. [in Russian with English summary]. 



i82 V. F. EASTOP 

Saemann, D. 1966. Beitrag zum Vorkommen und Massenwechsel auf Koniferen lebender 
Lachniden (Homoptera, Aphidina) im Erzgebirge wahrend des Jahres 1964. Hercynia 
3 : 374-386. 

Schels, J. 1957. Beobachtungen an Cinara (Lachnus) pinicola Kaltenbach, (Lachnus 
hyalinus Koch). Z. Bienenforsch. 4 : 151-179. 

Schimitschek, E. 1944- F or stinsekten der Turkei und ihre Umwelt. Prague, 371 pp. [Aphi- 
doidea pp. 258-264]. 

Schouteden, H. 1906. Catalogue des Aphides de Belgique. Mini. Soc. r. ent. Belg. 12 : 
189-246. 

Schrank, P. von. 1801. Fauna Boica 2 (i) : 1-374. Ingolstadt (aphids pp. 102-139). 

Shaposhnikov, G. Kh. 1964. In Bey-Bienko, G. Y. Classification Keys to the insects 
of the European part of the U.S.S.R. Aphidinea 1 : 489-616. Acad. Sci., Moscow & Lenin- 
grad [in Russian, English translation in 1967 by Israel Program for Scientific Translations 
Ltd., Jerusalem. Aphidinea pp. 616-799]. 

Shinji, O. 1941. Monograph of Japanese Aphids. 1215 pp. Tokyo [in Japanese]. 

Smith, C. F., Martorell, L. F. & Escolar, M. E. P. 1963. Aphididae of Puerto Rico. 
Tech. Pap. agric. Exp. Stn P. Rico 37 : 121 pp. 

Smith, R. C. 1932. A summary of the population of injurious insects in Kansas for 1931. 
/. Kans. ent. Soc. 5 : 65-92. 

Stenseth, C. & Bakke, A. 1968. Aphids of the Family Lachnidae found on Conifers in 
Norway. Meddr. norske SkogsforsVes. 89 : 233-238. 

Stroyan, H. L. G. 1955. Recent additions to the British Aphid Fauna. Part II. Trans. 
R. ent. Soc. Lond. 106 : 283-340. 

1957- Further additions to the British Aphid Fauna. Trans. R. ent. Soc. Lond. 109 : 

311-360. 

1964. Notes on hitherto unrecorded or overlooked British aphid species. Trans. R. 

ent. Soc. Lond. 116 : 29-72. 

Swain, A. F. 1919. A synopsis of the Aphididae of California. Univ. Calif. Pubis Ent. 

3 (1) : 1-221. 
1921. Miscellaneous Studies in the Family Aphidae. (Hem. Horn.) V. Notes on some 

Lachnids in the British Museum. Ent. News 32 : 209-213, 225-229. 
Szelegiewicz, H. 1962a. Materialmen zur Kenntnis der Blattlause (Homoptera, Aphididae) 

Bulgariens. Annls zool. Warsz. 20 : 47-65. 

1962b. The Identity of Lachnus nudus Mordwilko, 1895 (Homoptera, Aphidina). Bull. 

Acad. pol. Sci. CI. II Sir. Sci. biol. 10 : 21-22. 

1962c. Zur Validitatsfrage der Art Cinara pinihabitans (Mordv.) (Homoptera, Aphididae). 

Bull. Acad. pol. Sci. CI. II Sir. Sci. biol. 10 : 245-249. 

1962^. Contribution to the knowledge of Polish aphids (Homoptera, Aphididae) I. 

Subfamily Lachninae. Fragm. faun. 10 : 63-98 [in Polish with Russian & German 
summaries] . 

1963. Blattlause (Homoptera, Aphididae) aus der Mongolei. Annls zool. Warsz. 21 : 

109-142 + 7 plates. 

Takahashi, R. 192 1. Aphididae of Formosa. I. Spec. Rep. Formosa agric. Exp. Stn 20 : 
1-97. 

1931. Aphididae of Formosa. 6. Rep. Govt Res. Inst. Dep. Agric. Formosa 53 : 1-127. 

1937- Additions to the aphid fauna of Formosa (Hemiptera), IV. Philipp. J. Sci. 

63 : 1-19. 

1941. Some foreign Aphididae. Insecta matsum. 15 : 146-150. 

Taschev, D. G. 1 961. New plant lice (Horn., Aphid.) of the fauna of Bulgaria. God. sof. 
Univ. Biol. 53 : 157-160 [in Bulgarian]. 

1964. Contribution to the knowledge of the aphids (Horn., Aphid.) of Rosental. God. 

sof. Univ., Biol. 57 : 171-188 [in Bulgarian]. 

Theobald, F. V. 1914. African Aphididae. Bull. ent. Res. 4 : 313-337. 



REVIEW OF CINARA IN BRITAIN 183 

1926-1929. The Plant Lice or Aphididae of Great Britain 1 : 1-372 (1926); 2 : 1-411 

(1927); 3 : 1-364 (1929). Ashford, Kent. 

Thomas, I. 1948. Common Names of Aphididae. Entomologist's mon. Mag. 84 : 155-161. 

& Jacob, F. H. 1940. A List of Aphididae collected mainly in North Wales during 

1938 and 1939. NWest. Nat. 1940 : 139-153. 

Tissot, A. N. 1939. Notes on the Lachnini of Florida with Descriptions of Two New Species 
(Homoptera: Aphiidae). Fla Ent. 22 : 33-48. 

1945- Additions to the Lachnini of Florida (Homoptera: Aphididae). Fla Ent. 27 : 

43-54 (1944)- 
& Pepper, J. O. 1967. Two new species of Cinara (Homoptera: Aphididae) associated 

with Pine Rust Lesions. Fla Ent. 50 : 1-10. 
Toth, L. 1935. Beitrage zur Kenntnis der Aphidenspeicheldruse. Z. Morph. Okol. Tiere 

30 : 495-505- 
Tuatay, N. & Remaudiere, G. 1965. Premiere contribution au catalogue des Aphididae 

(Horn.) de la Turquie. Revue Path. vig. Ent. agric. Fr. 43 : 243-278. 
Wahlgren, E. 1939. Revision von Zetterstedt's lapplandischen Aphidina. Opusc. ent. 

4 : 1-9. 

Walker, F. 1848. Descriptions of Aphides. Ann. Mag. nat. Hist. (2) 2 : 43-48, 95-109, 

190-203, 421-431. 

1849. Descriptions of New British Aphides. Zoologist 7 appendix : xxxi-xl, xliii-lvii. 

1852. List of the Specimens of Homopterous Insects in the Collections of the British Museum, 

Part IV, London, 1188 pp. + 9 plates [Aphids pp. 934-1065]. 
Waterston, J. M. 1949. The Pests of Juniper in Bermuda. Trop. Agric, Trin. 26 : 5-15. 
Way, M. J. 1963. Mutualism between ants and honeydew-producing Homoptera. A. 

Rev. Ent. 8 : 307-344. 
Weed, C. M. 1890. Fourth Contribution to Life History of Little known Plant-Lice. Bull. 

Ohio agric. Exp. Sin 1 : 111-120. 
Weigel, C. & Baumhofer, L. G. 1948. Handbook on insect enemies of flowers and shrubs. 

Misc. Pubis U.S. Dep. Agric. 626 : 115 pp. 
Weis, S. 1955. Die Blattlause Oberosterreichs I. (Homoptera, Aphidoidea). Ost. zool. Z. 

5 : 464-559- 

Wellenstein, G. 1930. Beitrage zur Systematik und Biologie der Rindenlause (Lachninae 

CB.). Z. Morph. Okol. Tiere 17 : 737-767. 
Werder, O. 1931. Beitrage zur Kenntnis der Aphiden-Fauna von Basel und Umgebung. 

Verh. Naturf. Ges. Basel 42 : 1-98. 
Westwood, J. O. 1840. An introduction to the modern classification of insects, II. London, 

pp. 587 + 158 pp. of synopsis & addenda [Aphididae pp. 437-442]. 
Wilson, G. F. 1948. Two injurious Aphid Pests of Conifers. Jl R. hort. Soc. 73 : 73-78 + 

2 plates. 
Wilson, H. F. 191 i. Notes on the synonymy of the genera included in the tribe Lachnini. 

Ann. ent. Soc. Am. 4 : 51-54. 
1919. Some new lachnids of the genus Lachniella (Homoptera-Hemiptera). Can. Ent. 

51 : 18-22. 
& Vickery, R. A. 191 8. A species list of the Aphididae of the world and their recorded 

food plants. Trans Wisconsin Acad. Sci. Arts d>- Letters 19 : 22-355. 
Withycombe, C. L. 1923. Notes on the Biology of some British Neuroptera (Planipennia). 

Trans, ent. Soc. Lond. 1922 : 501-594. 
Wood-Baker, C. S. 1951. Records of five European aphids (Hem.). Entomologist's, mon. 

Mag. 87 : 271. 
Yakhontov, V. V. 1929. A list of Pests injurious to economic Plants of Boukhara District 

and their Parasites and Predatory Insects. Trudy shirabud. opyt. sel'.-khoz. Sta. 2 : 1-46 

[in Russian]. 
Zeck, E. H. 1948. The Pine Aphid. Agric. Gaz. N.S.W. 59 : 422, 426. 



i8 4 V. F. EASTOP 

Zetterstedt, J. W. 1828. Fauna Insectorum Lapponica. Hamm, 563 pp. [Aphidiae pp. 

550-559]- 
1838-1840. Insecta Lapponica. Leipzig 1140 pp. 1-687 (1838); 868-1013 ( J 839); 

1014-1140 (1840) [Aphididiae pp. 306-313]. 
Zimmerman, E. C. 1948. Insects of Hawaii 5, Homoptera: Sternorhyncha. Honolulu, 

464 pp. [Aphididae pp. 53-131]. 
Zirnits, J. 1927. Beitrage zur Kenntnis der Aphiden Lettlands. Z. wiss. InsektBiol. 

22 : 244-256. 
Zoebelein, G. 1956. Der Honigtau als Nahrung der Insekten. Z. angew. Ent. 38 : 369-416 

& 39 : 129-167. 



INDEX 



abamaleki, 160, 163 

abieticola, 105, 106, 113, 116, 117, 123-128, 

150-152 
abietis Matsumura, 104, 105, 149 
abietis Walker, 132, 153 
acutirostris, 109, 112, 117, 127-129, 139 
agilis, 105 
alacra, 149 
Aphis, 104, 123, 129, 132, 134, 141, 143, 146, 

148, 150, 152, 153, 156, 160, 162, 163 

biotae, 167 

boerneri, 105, 106, 108, 109, 112, 113, 118, 

129-131, 147 
bogdanowi, 105, 106, 108, 113, 114, 117, 120, 

128, 132-133 
borealis, 123, 132 
braggii, 156 

brauni, 105, 106, no, 118, 119, 133-134 
brevispinosa, 162 

Buchneria, 105, 106, 108, 148, 149 
bulgarica, 125, 126 

callitris, 167 

canadensis, 167 

canatra, 139, 162 

cecconii, 105, 106, 123, 125, 126, 128 

chinookiana, 106 

cilicica, 105, 106, 123, 125, 128 

Cinara, 103-170 

Cinarella Borner, 105, 129 

Cinarella Hille Ris Lambers, 105-107, 156, 

164 
Cinarellia, 105, 129 
Cinaria, 105, 106, 125, 128, 132, 139, 144, 

146, 150, 160, 161 
Cinarina, 105, 106 
Cinaropsis, 105-108, 132, 134, 136, 150, 153, 

165 



cistatus, 134, 165, 166 

coloradensis, 107 

comata, 106 

confinis, 123, 125, 126 

costata Zetterstedt, 104, 106, 114, 121, 

134-136, 166 
costatus Hartig, 134 
cuneomaculata, 129, 131, 143, 146 
cupressi, 109, 112, 115, 122, 136-140, 170 
Cupressobium, 103, 105-108, 121, 122, 137, 

140, 167 
curvipes, 152 

difficilis, 106, 139 

Dilachnus, 105, 106, 123, 125, 132, 137, 141, 

146, 148, 150, 153, 156, 163, 165, 167 
Dinolachnus, 105, 106, 125, 128 
doncasteri, 146 
drastichi, 165 
Dryobius, 134 

escherichi, 105, 106, no, 112, 117, 139, 162 
Eulachnus, 105, 133, 148, 156, 160 
excelsae, 160 
ezoana, 133 

farinosus, 134 

fasciatus, 104, 106, 134, 136 

flavus, 153 

fornacula, 149 

fresai, 112, 115, 123, 138, 140-141 

gracilis, 104-106 
grande, 128 

grossus, 123, 125, 128, 150 
guadarramae, 160, 162 

hattorii, 127 
horii, 166 



INDEX 



185 



hottesi, 107 
hyalinus, 153, 156 

intermedia, 132 

juniperi, 105, 106, 115, 123, 136, 137, 

140-143, 167, 170 
juniperinus, 136-138, 140, 141 

kochi, 144, 146 

kochiana, 105, 106, 109, no, 116, 131, 

143-146 
konoi, 106, 127 
kosarowii, 164 

Lachniella, 104, 106, 108, 123, 129, 134, 136, 
139, 141, 143, 146, 148, 153, 156, 160, 163, 
166 

Lachnus, 104, 123, 125, 129, 132, 134, 136, 

137. J 39, 14 1 . H3. I4 6 - M8. 150. i5 2 . 153. 

156, 160, 163, 165, 166, 167 
Laricaria, 105, 106, 144 
laricicola Borner, 105, 106, 129 
laricicola Matsumura, 105, 106, 129, 131, 148 
laricifex, 131, 146, 148 
laricifoliae, 148 
laricis Hartig, 145-148 
laricis, Koch, 129, 143-146 
laricis Walker, 105, 106, 108, no, 116, 117, 

120, 129, 132, 143, 146-148, 150, 156 
lasiocarpae, 106, 125, 126 
longipennis, 106, 127 
longipes, 150 
longirostvis, 160, 163 
lyallii, 148 

macchiatii, 148 

macrocephalus, 153 

maculosus, 146 

maghrebica, 163 

maidis, 159 

manitobensis, 107 

maui, 140 

Mecinaria, 105, 106, 132, 150, 152 

mediterraneum, 167, 170 

mingazzinii, 105 

minor, 165 

montanicola, 128, 160, 162 

mordwilkoi, 140, 141, 143 

muravensis, 146 

Neochmosis, 105, 106, 132, 137, 141, 146, 148, 

153. 167 
Neocinara, 105, 106 



Neodimosis, 105, 163 
neubergi, 105, 106 
nigrofasciatus , 133 
nigrotubercalata, 146 
nimbata, 106, 136 
nopporensis, 153 
nudus, 139, 160-162 

palaestinensis, 162 

Panimerns, 105, 106, 123, 129, 134, 137, 146, 

148, 150, 153, 156, 160, 163, 167 
pasheki, 150 
pectinatae, 105, 106, 109, no, 117, 119-121, 

148-149 
piceae, 105, 106, 108-110, 112, 116, 119, 123, 

125, 128, 148, 150-152 
piceicola, 132, 153, 156, 165-166 
pichtae, 148 

picta, 139, 160 

pilicornis, 105, 106, 108, 113, 114, 121, 122, 

126, 152-156, 160 
pilosa, 156, 160, 162 
pinata, 144, 162 

pinea, in, 117-119, 156-162 

pineti Fabricius, 152, 156, 160 

pineti Hartig, 156 

pineus, 103, 106, 156 

pini, 104, 106-108, 112, 116, 139, 156, 

160-162 
pinicolus, 105, 106, 129, 146, 152, 153, 156, 

160, 163 
piniformosana, 159 
pinihabitans, no, 120, 121, 134, 153, 160, 

162-163 
piniphila, 105 
piniradicis, 144 
Pitsaria, 125, 128 
Pityaria, 105, 132 
poly seta, 160 
Protolachnns , 105 
pruinosus, 105, 106, 132 
Pseudocinara, 105, 106 
Pterochlorns, 134 
pubescens, 125, 126 
puerca, 144 

radicicolus, 132, 133 
Rhopalosiphum, 159 
roboris, 136 

sabinae, 137, 138 

schimitscheki, 113, 114, 116, 118, 164-165 

Schizoneura, 134 

setosa, 160, 162 



1 86 



INDEX 



signata, 136 

stichensis, 136 

sonata, 106, 128 

spiculosa, 148 

stroyani, 112, 114, 122, 156, 165-166 

Subcinara, 105, 106, 133 

symphiti, 134 

taedae, 162 

taeniata, 129, 156, 160, 163 

tanneri, 144 

taxicola, 165 

tenuior, 146 

thatcheri, 144, 162 

thujae, 137 



thujafilinus, 137 

thujaphilinum, 166, 167 

thujafolia, 166, 167 

thujafoliae, 167 

todocolus, 106 

Todolachnus, 104, 105, 125, 149 

tujae, 136, 137 

tujafilina, 112, 115, 122, 123, 141, 166-170 

vanduzei, 123, 125, 126, 128, 150 
viridescens, 105, 106, 132 

wacassasae, 140 
Wilsonia, 104, 106 
winonkae, 167 




V. F. Eastop, D.Sc, 

Department of Entomology 

British Museum (Natural History) 

Cromwell Road 

London, SW7 5BD 



A LIST OF SUPPLEMENTS 

TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera) . Pp. 177: 
18 plates, 270 text-figures. August, 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 : 500 text-figures. September, 

1965. £3-25- 

5. Ahmad, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156: 
475 text-figures. November, 1965. (out of print) £2.15. 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. 

£3.15. 

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 
text-figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera). Pp. 509. £8.50. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322 : 348 text-figures. August, 1967. £8. 

11. Mound, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp.172: 
82 text-figures. May, 1968. £4. 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. 
November, 1968. £$. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
Nymphalidae). Pp. 155 : 3 plates, 101 text-figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North-Western Europe 
(Hymenoptera : Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 
68 plates, 15 text-figures. October, 1971. £12. 



»(««£ Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 



THE SIMULIIDAE DESCRIBED BY 
N. BARANOV AND THEIR TYPES 

(DIPTERA) 



R. W. CROSSKEY 

AND 

B. V. PETERSON 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 27 No. 3 

LONDON : 1972 



THE SIMULIIDAE DESCRIBED BY N. BARANOV 
AND THEIR TYPES (DIPTERA) 




BY 

ROGER WARD CROSSKEY 

Commonwealth Institute of Entomology 
AND 

BOB VERN PETERSON 

Entomology Research Institute, Canada Department of Agriculture 



Pp. 187-214 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 27 No. 3 

LONDON : 1972 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 27, No. 3 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. not. Hist. (Ent. 



© Trustees of the British Museum (Natural History) 197^ 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 13 April, 1972 Price £1.05 



THE SIMULIIDAE DESCRIBED BY N. BARANOV 
AND THEIR TYPES (DIPTERA) 

By R. W. CROSSKEY & B. V. PETERSON 

CONTENTS 

Page 
Synopsis ........... 189 

Introduction .......... 189 

Acknowledgements ......... 194 

Short biography of N. Baranov ....... 194 

Baranov's nominal species-group taxa of Simuliidae and their 

types ............ 196 

Bibliography . . . . . . . . . . 211 

Index to species-group names . . . . . . .214 

SYNOPSIS 

An alphabetical catalogue is given of the 43 species and infraspecific taxa of Simuliidae 
described by N. Baranov, with an account of all existing type-material on which the names are 
based. Sixteen lectotypes are designated. Each species-group taxon is assigned to its correct 
genus-group segregate after type-examination, and previously established synonymies are 
given; no new synonyms are involved. A complete bibliography of Baranov's works on 
Simuliidae is included, although most of them are not taxonomic. 

INTRODUCTION 

The extensive taxonomic and other investigations on the European Simuliidae 
made in the past 25 years, since the last war, have taken little account of the work of 
Mr Nikola Baranov between 1924 and 1942 on the black-fly fauna of Yugoslavia, 
and even Rubzov's (1959-1964) comprehensive monograph of the Palaearctic 
Simuliidae mentions only nine of the 43 species-group taxa that Baranov described. 
It is quite likely that some of Baranov's names apply to, and have priority over, 
taxa and names that have been proposed recently by other authors, though it will 
be some time before all the various synonymies among European black-flies can be 
unmasked. 

The European literature of simuliidology is bedevilled with nomenclatural diffi- 
culties arising from too much neglect of the early names, misidentification through 
failure to study the types, and a lack of attention on the part of too many workers 
to the elements of nomenclature. Before many of the problems of identity and 
synonymy can be resolved it will be necessary to make extensive studies of the 
types, and a preliminary need is for an up-to-date catalogue of Palaearctic black- 
flies which includes the basic data on all the primary types and their whereabouts. 
This is being prepared at the moment by one of us (R.W.C.), and the present con- 
joint paper on Baranov's Simuliidae has two main purposes: to make known to 
other simuliidologists what material still exists in Baranov's collection and to clarify 
a number of matters concerning Baranov's Simuliidae in advance of the projected 
catalogue. 



igo R. W. CROSSKEY & B. V. PETERSON 

Most of Baranov's work in Yugoslavia during the 1920's and 1930's was concerned 
with the infamous 'Golubatz fly', Simulium columbaczense (Schonbauer) [name 
now a synonym of S. colombaschense (Fabricius)], which breeds along a stretch of 
the river Danube and becomes an appalling and notorious biting pest in periodical 
outbreaks, and Baranov himself well documented his extensive studies on this 
pest (see references). It was as a by-product from this main work that Baranov 
discovered and named the various species, subspecies, forms and races of Simulium 
Latreille s.l. (numbering 43 taxa in all) that he described; all were found in Yugo- 
slavia and most were described from the states of Serbia and Macedonia. 

Baranov made a personal collection of Yugoslav Simuliidae which consisted 
both of pinned adult flies (many of them reared from pupae and kept with their 
associated pupal pelts) and of adults and early stages preserved in alcohol, and this 
collection originally contained all the type-specimens on which his taxa were based. 
Unfortunately part of the collection, including all of the alcohol material, was lost 
at the end of the second world war, when Baranov was experiencing great difficulties 
as a displaced person, and has never been found again. (Our information on this 
point was obtained from Mr Baranov himself, both from conversations during the 
preparation of this paper and from a letter that he wrote to Mr Guy Shewell, of the 
Division of Entomology, Canada Department of Agriculture, in 1954 when the 
purchase of the remainder of his collection was being negotiated.) 

The surviving part of Baranov's collection consists of adult material, often with 
associated pupal pelts, and is in the Canadian National Collection (hereafter abbre- 
viated CNC). It was bought from Baranov in 1954 by the Canada Department of 
Agriculture and is now housed as part of the CNC in the Entomology Research 
Institute at Ottawa, Canada. All extant types of Baranov's taxa of Simuliidae are 
in this collection except for some paralectotypes deposited in the British Museum 
(Natural History) (text-abbreviation: BMNH) by an exchange made whilst this 
paper was in preparation. Baranov himself did not exchange specimens with other 
workers or deposit 'cotypes' in other museums (Baranov, pers. comm., and i. litt. 
to Shewell as follows: 'Your assumption that in no other collection [besides his 
own] these cotypes are contained is correct. Since I had to stop my work before I 
had finished I never started to exchange material with anybody in this line 
[Simuliidology]'). 

Baranov's letter to Shewell, referred to above and dated 4th March 1954, contains 
some helpful and definite information on the fate of several of the types that are 
lost, and we have accordingly cited this letter from time to time in the body of this 
paper: the text notation 'Baranov i. litt. to Shewell' refers always to this particular 
letter. 

Baranov described 43 nominal taxa of species-group Simuliidae, and five genus- 
group taxa. The latter, viz. Danubiosimulium, Echino simulium, Pseudodagmia, 
Pseudonevermannia and Pseudo simulium, are discussed by Stone (1963) and are not 
considered further in this paper. Baranov's collection in Ottawa contains specimens 
labelled as primary types of seven species-group entities, in addition to the still 
extant types of the described taxa, but we are now convinced that the seven names 
attached to these extra 'types' were never actually published (since they are 



BARANOV'S TYPES OF SIMULIIDAE 191 

manuscript names we are not recording them in print). Baranov (i. litt. to Shewell), 
when asked about these names in 1954, implied that they had been published al- 
though he was not able to recall the references, and because of the possibility that 
they could have appeared in print in one of the little-known Yugoslav journals 
and been overlooked we have made extra-careful searches for them. No trace of 
any of these seven names has been found from a page-by-page search through all 
the works on Simuliidae of which Baranov was sole or conjoint author, and our 
bibliography of these works is, we believe, complete ; we conclude, and Mr Baranov 
(personal communication with R.W.C.) now thinks this must be so, that Baranov 
had intended to publish the seven names but never actually did so (either because 
no manuscripts on the seven taxa were completed or because they went astray 
after completion, perhaps during the difficult war years). Of the 43 published taxa 
it should be explained that, for reasons given where appropriate in the text, we 
regard Odagmia ruficornis and 0. ruficornis prima as names applying to a single 
nominal taxon, and likewise Wilhelmia stylata and W. stylata prima as likewise 
applying to one taxon only; three of Baranov's names, viz. barensis, intermedia 
and mazedonica were overlooked by Smart (1945) and omitted from his catalogue 
of world Simuliidae. 

When originally described Baronov's 43 taxa comprised 11 species, six subspecies, 
24 forms (plus two uses of f. prima for the nominate subdivision of species ruficornis 
and stylata) and two ecological races. Some of these taxa were well and formally 
described, but for others there is only the bare minimum of descriptive matter — 
often not set out as a formal description — to make the names nomenclaturally 
available. Nevertheless all of them have, in our opinion, status in nomenclature 
(none can be clearly interpreted as infrasubspecific in the meaning of the International 
Code of Zoological Nomenclature, 1961, for the reasons given below), and are available 
names (though some, as annotated in the text, are primary homonyms or secondarily 
homonymous in Simulium s.L). Baranov himself fixed types for all his taxa, and 
intended that his various forms and races should have status in nomenclature as 
much as his subspecies and species. He divided species either into subspecies or 
into forms (or in the case of S. columbaczense into geographically separate ecological 
races), but his subspecies were never themselves divided into forms, and there is 
no instance in which any of his forms is infrasubspecific; furthermore the form 
names were bestowed on geographically allopatric entities (see, for instance, Bara- 
nov's 1937a segregation of forms in the 'Vardar-Formenkomplex' and the 'Morava- 
Donau-Formenkomplex') not different in kind from his subspecies, and indeed he 
sometimes referred to his subspecies as forms without discrimination. Hence all 
of his form names, as none is clearly infrasubspecific, have nomenclatural status 
under the Code and are based upon the types. Similarly the two entities originally 
described as ecological races (litorale and profundale) are here deemed to have status 
under the Code since these names cannot be interpreted as infrasubspecific (because 
the specific taxon to which they belong was not divided into subspecies, because 
Baranov himself variously referred to them as races or forms, and because they 
were described as geographically as well as ecologically separated taxa). 

One other matter concerning Baranov's form names requires mentioning. 



192 R. W. CROSSKEY & B. V. PETERSON 

Baranov used each of the names prima and secunda five times for supposedly distinct 
forms within five species, all of the uses of these names being published in a single 
paper (Baranov, 19266). In practice the existence of five prima homonyms and 
five secunda homonyms creates no difficulty, because (with the exception of one use 
by Rubzov, 1962) later authors have not regarded Baranov's prima and secunda 
entities as having taxonomic validity; Smart (1945) merely listed them under the 
appropriate species name. In the present paper we draw attention to the homonymy 
and point out that future authors may need to provide replacement names in the 
improbable event that they should consider some of Baranov's prima and secunda 
forms worthy of named rank. There is as yet no determined senior homonym 
among the uses of prima, but Rubzov (1962 : 412) has given Wilhelmia equina 
secunda Baranov full specific rank and we consider that Rubzov's citation determines 
which of the five uses of secunda is to be held as the senior homonym. 

In the body of this paper we enumerate all of Baranov's described species-group 
taxa in alphabetical order of their original combinations and give details of the 
types. For each name the entry is arranged to show the following information in 
the order indicated :- 

Name; author; date and page reference of original publication; status and 
sex of primary type (or syntypes) ; present lectot}'pe designation (if necessary) ; 
locality and date of primary type ; type-depository. 

Number and sex of paralectotypes or paratypes, with data and depository 
information as for primary types. 

Explanatory comments or annotations when necessary. 

Genus-group assignment for each name where possible, and annotations on 
correct placement. 

For the localities we have always cited a larger unit first, i.e. state before town 
or village, town before stream name, and we have always indicated the state (e.g. 
Serbia or Macedonia) for each locality even though Baranov often omitted this 
information from the original publication. Baranov, in publication and on labels, 
used the spelling Skoplje for one of his important localities, but we have thought it 
better to use the English spelling Skopje. Baranov variously used either the 
Croatian spelling Golubac or the German spelling Golubaz; we have consistently 
used the former spelling, which accords with English transliteration from Serbian. 

Whilst working through Baranov's collection in conjunction with the publications 
containing his descriptions we have found a number of anomalies. Most of the 
specimens in the collection bear Baranov's labels either as 'Typus' or as 'Cotypus', 
but there is not always complete agreement between the status indicated by Baranov 
and that which is apparent from the description; there are some discrepancies 
between the sex of the specimen labelled as type and that described in publication, 
and there are several instances in which specimens labelled as types by Baranov 
cannot have any type-status because they were not collected (as indicated by the 
data labels) until some time after the description had been published. We have 
judged each case of discrepancy individually, and have determined the status of 
types in doubtful cases after careful scrutiny of the original publications; in some 



BARANOV'S TYPES OF SIMULIIDAE 193 

instances it has seemed reasonable to accept certain specimens as being original 
syntypes in the light of some piece of information contained somewhere in Baranov's 
text, but in other instances it has been considered that the evidence does not 
warrant inclusion of certain specimens in the syntypic series (e.g. too great a conflict 
between cited dates or localities or sex taken in combination). Any discrepancies 
between specimens and descriptions of the kind mentioned have been fully annotated. 
In the case of syntypic series we have ordinarily labelled and designated the specimen 
labelled 'Typus' by Baranov as the lectotype, provided there is no discrepancy (no 
lectotype designations have previously been published for Baranov's simuliid taxa 
and all lectotypes mentioned are newly designated). 

Baranov published several figures of the male hypopygium of various Simuliidae, 
including some of his new taxa, but all of his slides are believed lost ; there are none 
with the main collection in CNC. 

Baranov's species-group taxa of Simuliidae were described in nine taxonomic 
papers, but Baranov published many other papers on the Simuliidae of which some 
contain keys and other taxonomic information. No complete bibliography of 
Baranov's works on black-flies has up to now been published, even Rubzov's 
(1959-1964) monograph including little more than half the relevant references, so 
we have thought it useful to bring together such a bibliography in the present paper. 
Our attempts to trace the manuscript names (already referred to) in print obliged 
us to examine every one of Baranov's works on the Simuliidae, many in obscure 
publications overlooked by recording and abstracting journals, and rather than 
'waste' the bibliographic information obtained we have assembled it as a full 
bibliography in the references at the end of this paper. To the best of our know- 
ledge this bibliography is complete. 

The bibliography contains in all 34 publications of Baranov concerned with 
Simuliidae (30 in which he was sole author and 4 in which he was co-author). 
Out of this total of works only n titles are to be found in the Zoological Record, 
and only 20 are recorded in the Review of Applied Entomology, Series B, Medical & 
Veterinary (17 with abstracts and three by title only). This does not, however, 
reflect badly on these recording publications, for two reasons : firstly, several of the 
publications are in the nature of instruction pamphlets or semi-popular accounts 
of the Golubatz fly (not strictly scientific papers), and, secondly, many of the papers 
were published in local Yugoslav journals which are either not held in any British 
library or are held in incomplete sets (e.g. lacking volumes for the second world 
war years) . Although so many of the papers are in rather inaccessible publications 
it has been possible for one of us (R.W.C.) to see thirty of them in the original, and 
the remaining four in photocopy, by bringing copies together from the following 
sources: personal gift and loan from Mr Baranov himself of a few publications still 
remaining in his possession; library holdings of British Museum (Natural History), 
Commonwealth Institute of Entomology (which holds many reprints, including 
most of the important papers published in Veterinarski Arhiv), and of the Common- 
wealth Bureau of Animal Health, Weybridge ; and photocopies from the Nacionalna 
i Sveucilisna Biblioteka, Zagreb, the library of the Veterinary Faculty, Institute 
of Medical Researches, Belgrade, and the library of the U.S. Department of Agricul- 



194 R- W. CROSSKEY & B. V. PETERSON 

ture, Washington, D.C. References from all sources have been cross-checked against 
a manuscript of his simuliid papers prepared by Baranov himself in the 1950's 
(copy supplied to us by Dr Alan Stone), and with Mr Baranov's recollections at the 
present time of his various publications. The manuscript list referred to was 
prepared by Mr Baranov largely from recollection, when he was living in Pakistan 
without access to literature, and as a result there are a few minor omissions and 
errors; nevertheless it has been of such immense value to us in providing leads to 
many of the lesser known works (which are not cited in the recording journals and 
could easily have been overlooked) that we take this opportunity of recording our 
indebtedness to this list. 

Finally in this introduction we should note that we have consistently used the 
spelling Baranov with terminal V throughout the paper, though Baranov some- 
times used the alternative 'ff' ending to his name (especially in German-language 
publications). 

ACKNOWLEDGEMENTS 

It gives us great pleasure to acknowledge the generous help that we have received 
from Mr Nikola Baranov, including valuable information on his work in Yugoslavia 
and on his simuliid collections and publications, and the generous gift or loan of 
copies of some of his works. One of us (R.W.C.) is also most grateful to Mr and Mrs 
Baranov for their kindness and hospitality during a visit (in January, 1971) to their 
home in Shepherd's Bush, West London, when many of the points that had arisen 
in the preparation of this paper were helpfully discussed. 

We are grateful also to the following persons who helped us by providing photo- 
copies of various publications: Mr Milutin Ivanusic, National and University 
Library, Zagreb; Dr Alan Stone, U.S. Department of Agriculture, Washington, 
D.C. ; and Dr Vera Zivkovic, Veterinary Faculty, Institute of Medical Researches, 
Belgrade. We thank Dr Zivkovic also for helpful information on Baranov's publica- 
tions and for confirming that no specimens from Baranov's material are in Yugoslav 
collections. Dr Curtis Sabrosky very kindly supplied us with information on 
Baranov's son which enabled us, in turn, to establish contact with Mr and Mrs 
Baranov. Miss Pamela Gilbert, of the Department of Entomology Library, 
British Museum (Natural History), helped us considerably with the unusually obscure 
references involved in this work, and in obtaining photocopies from other libraries, 
and for this we are grateful. 

SHORT BIOGRAPHY OF N. BARANOV 

Nikola Baranov was born at Orel in Russia in 1887. He grew up in Nishni 
Novgorod, where his father was director of the grammar school, and graduated from 
the Natural History Faculty of the University of Moscow. He then specialized in 
applied entomology, studying under Professor Kulagin at the Agricultural Academy 
in Petrovsko Rasumovsko (near Moscow), and — after completing the course in 
applied entomology — was appointed entomologist to the Agricultural Station at 
Kursk in southern Russia. Here he was working as an agricultural entomologist 
at the time of the Russian revolution in 1917. 






BARANOV'S TYPES OF SIMULIIDAE 195 

In the autumn of 1919 Baranov was forced to leave Kursk because of the civil 
war. He went first to the Crimea, but decided to leave Russia finally from there in 
the autumn of 1920. He settled in Yugoslavia, and in May 1921 was appointed to 
the staff of the Agricultural Station (Poljoprivredne Stanica) at Topcider, Belgrade, 
where he started entomological work on a variety of agricultural pest problems. 
He soon began to work in particular on the Acridoidea, especially the Moroccan 
locust pest Dociostaurus maroccanus (Thunberg). His first scientific papers appeared 
in 1924 and 1925 and were concerned with the morphological variability and natural 
enemies of this locust, and with the classification of Serbian Acridoidea by using the 
male genitalia. As a result of studying the dipterous parasites of Dociostaurus 
he acquired an interest in Sarcophagidae and Tachinidae, of which (in later years) 
he described many new genera and species (see Sabrosky & Crosskey, 1969, 1970). 

After five years at the Topcider station Baranov took the post of entomologist 
at the Institute of Hygiene in Skopje (Macedonia) in 1926, from where he was 
shortly transferred to the Institute of Hygiene at Zagreb (Croatia) when this was 
newly opened in 1928. In the following year a lady assistant from Vienna was 
appointed to help Baranov with his entomological work, and they were married in 
1931 ; from this time on Baranov was greatly aided in his work by Mrs Baranov, 
and it is of special interest to note that it was she who drew nearly all the figures in 
his papers and put into German those papers which he published in this language. 
Baranov began to work upon the Simuliidac whilst still at Topcider because of the 
great importance as a livestock pest of the Golubatz fly (Simulium colombaschense 
(Fabricius), syn. columbaczense Schonbauer), and later whilst at the Institutes of 
Hygiene at Skopje and Zagreb he extended his interests to the whole Yugoslav 
simuliid fauna; during the 1930's the Veterinary Faculty at Zagreb supported his 
well documented studies on the Golubatz fly (for further information see Introduc- 
tion). Baranov's duties with the Institutes of Hygiene were many and varied and 
the Golubatz fly was only one aspect; in addition he worked on domestic flies of 
hygienic importance, including the control of house-flies and malarial mosquitoes. 
For many years, in conjunction with the Agricultural Station in Zagreb, he helped 
to organize the control measures against insect pests of olive trees in the coastal 
areas of Dalmatia. 

Baranov continued to work with the Institute of Hygiene at Zagreb until 1944, 
when he left Yugoslavia and his entomological work ended. Immediately after 
the second world war Baranov was, for a time, a displaced person, but luckily 
throughout this difficult period he was able to keep most of his entomological 
collections intact, and to take them with him in 1948 to the newly-formed state of 
Pakistan (where Mrs Baranov was offered employment and Baranov was able to 
settle). The collections were later sold, the Simuliidae (plus some Culicidae and 
Muscidae) going to the Canada Department of Agriculture, Ottawa in 1954, and the 
Tachinidae and Sarcophagidae to the United States National Museum, Washington, 
D.C. in i960. 

In 1962 Baranov came to England, where — now aged 84 — he is living in retire- 
ment in London. 

We take this opportunity of thanking Mr and Mrs Baranov for providing much 



ig6 R. W. CROSSKEY & B. V. PETERSON 

of the information for this short account and of extending our best wishes to them 
for the future. 



BARANOV'S NOMINAL SPECIES-GROUP TAXA OF SIMULIIDAE AND THEIR TYPES 

Danubiosimulium columbaczense (Schoenbauer), form intermedia Baranov, 1939a : no, 
118-120 (Croatian), 122-125 (German). Syntypes o\ 9. pupae, larvae, Yugoslavia: Serbia, 
R. Danube, near Sip Canal, iv. 1938 (not located, presumed lost). 

Baranov proposed the name intermedia for a form of columbaczense (Schoenbauer) in which 
the adult flies were intermediate between those of columbaczense race litorale Baranov and 
columbaczense race profundale (being large like the former but having a metallic sheen like 
the latter) and emerged from the Danube in the geographical area of the Sip Canal earlier 
in the year than race litorale. Apart from adults Baranov mentioned larvae and pupae. 
The work containing the name intermedia has scattered mentions of characteristics differen- 
tiating the taxon, although there is no formal description, and the name is nomenclaturally 
available. No original type-material or any other material pertaining to intermedia has 
been located and the syntypic specimens are presumed lost. 

The name intermedia appears in the Rev, app. Ent. (B) 27 : 85-86 but was overlooked by 
Smart (1945) and omitted from his catalogue of world Simuliidae. It does not appear in 
Rubzov (1959-1964). Probably no present day taxonomist would regard intermedia as a 
taxonomic entity requiring a formal name and the name is best regarded simply as a synonym 
of Simulium columbaczense (Schoenbauer) ( = S. colombaschense (Fabricius)). It is a junior 
secondary homonym in Simulium s.l. of S. intermedium Roubaud, 1906, and would in any 
case, therefore, require a replacement name if applied to a valid taxon. 

Echinosimulium echinatum Baranov, 1938a : 313 (Croatian), 323 (German). Holotype 
pupa, Yugoslavia: R. Danube (not located and presumed lost). 

Baranov (1938a) described the larva and the pupa of this species and stated (Croatian text, 
p. 316, German text, p. 325) that the pupal description was based on the holotype. It is 
known that Baranov made slides of larval specimens for the larval description but these are 
not with the Baronov collection in CNC and are presumed lost; if later located, the larvae 
will have paratype status. 

Rubzov (1962 : 259) places echinatum in the segregate Titanoptevyx Enderlein { = Byssodon 
Enderlein) and this placement is undoubtedly correct, though whether subspecific status 
within maculatum Meigen is justified appears doubtful. 

Nevermannia angustitarsis (Lundstrom), form mazedonica Baranov, 1926b : 185, 193. 
Syntypes $, Yugoslavia: Macedonia, Hanrijevo near Skopje; Serbia, Tuman (lost). 

The type-material of this form was preserved in alcohol and was lost in Vienna at the end 
of the war (Baranov i.litt. to Shewell). A $ specimen from Skopje, 4.XL1926, is in Baranov's 
collection but has no type-status, as it was collected after publication of the name. 

The name remains enigmatic. Smart (1945) omitted the name mazedonica from his world 
catalogue of Simuliidae by oversight. 

Nevermannia aurea (Fries), form prima Baranov, 19266 : 185, 192. Syntypes S, ?» pupae, 
larvae, Yugoslavia: Macedonia, Hanrijevo near Skopje (lost). 

The type-material of this form was preserved in alcohol and was lost in Vienna at the end 
of the war (Baranov i.litt. to Shewell). 

The name remains enigmatic and is a secondary homonym in Simulium s.l. of prima 
Baranov (19266, forms of equina L., ornata Meigen, ruficornis Baranov and stylata Baranov). 



BARANOV'S TYPES OF SIMULIIDAE 197 

Nevermannia aurea (Fries), form secunda Baranov, 19266 : 185, 192. Syntypes c?, Yugo- 
slavia: Macedonia, Hanrijevo near Skopje (lost). 

The type-material of this form was preserved in alcohol and was lost in Vienna at the 
end of the war (Baranov i.litt. to Shewell) ; the original publication cited only the adult male, 
but it is possible that Baranov also had the female and immature stages at the time of 
description (as he did for aurea f. prima, see above). The name remains enigmatic and is a 
secondary homonym in Simnlium s.l. of secunda Baranov (19266, forms of equina L., ornata 
Meigen, ruficornis Baranov and stylata Baranov). 

Odagmia croatica Baranov, 1937a : 263 (Croatian), 274 (German). LECTOTYPE 9, by 
present designation, Yugoslavia: Croatia, Zagreb, 23.iii.1931 (CNC). 

Paralectotypes: 1 <J, 3 ?, same data as lectotype (J, 2 ? in CNC; one ? in BMNH). 

Baranov did not mention the number of specimens in the Croatian description (pp. 263- 
264), but mentioned one male and four females at end of the German description (p. 275); 
the type-specimens cited above therefore comprise the whole type-series. The paralectotypes 
are in poor condition and the male is on the same mount as one of the females. 

O. croatica is not placed by Rubzov (1959-1964); it is here confirmed from lectotype 
examination that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 503) gives the publication date of this name as 1936 in error. 

Odagmia croatica Baranov, form decolorata Baranov, 1937a : 264 (Croatian), 276 (German). 
LECTOTYPE $, by present designation, Yugoslavia: Croatia, Zagreb, 30.iii.1931 (CNC). 

Paralectotypes : 3 <J, 3 $, same data as lectotype (2 <J, 2 ? in CNC; 1 cJ, 1 ? in BMNH). 

Baranov mentioned both sexes in the original descriptions but did not state the number of 
specimens; all the above-listed specimens are considered to be original syntypes as their 
data fully conforms with the original publication. 

O. croatica f. decolorata is not placed by Rubzov (1959-1964); we confirm from lectotype 
examination that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 503) gives the publication date of this name as 1936 in error. 

Odagmia croatica Baranov, form nigrina Baranov, 1937a : 264 (Croatian), 275 (German). 
LECTOTYPE $, by present designation, Yugoslavia: Croatia, Zagreb, 23.iii.1931 (CNC). 

Paralectotypes: 2 $, 2 $, same data as lectotype (one of each sex in CNC and BMNH). 

Baranov did not mention the number of specimens in the Croatian description (p. 264), 
but mentioned two males and three females at the end of the German description (p. 276) ; 
the type-specimens cited above therefore comprise the whole type-series. All of them lack 
the abdomen. 

O. croatica i. nigrina is not placed by Rubzov (1959-1964); we confirm from lectotype 
examination that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 509) gives the publication date of this name as 1936 in error. 

Odagmia (Pseudodagmia) kondici Baranov, 1926a : 161. LECTOTYPE $, by present 
designation, Yugoslavia: Serbia, Tuman, 9.VL1925 (CNC). 

Paralectotypes: 3 $, same data as lectotype, on same mount (CNC). 1 pupal exuvium, 
Yugoslavia: Serbia, Golubac, 27. v. 1925 (CNC). 

Baranov described this species from both sexes, pupa and larva. We have not located 
any male or larval syntypes and presume these to be lost. The CNC collection contains 
(from Baranov's collection) specimens of kondici from Macedonia (2 cJ and 6 $ from Skopje 
and one $ from Treska) but these specimens have no type-status; they were collected in 
November, 1926 (Treska specimen) and July, 1927 (Skopje specimens), after the original 
description had been published. 

This species is assigned by Rubzov (1963 : 506), under the mis-spelling condici, to Teti- 
simulium Rubzov, and we confirm this placement after examination of the lectotype. 



198 R. W. CROSSKEY & B. V. PETERSON 

Odagmia ornata (Meigen), form anderliceki Baranov, 1937a : 263 (Croatian), 274 (German). 
Holotype $, Yugoslavia: Serbia, Golubac, Dedinje stream, 24.V.1936 (CNC). 

Paratypes: 1 $, 6 $, same data as holotype (CNC). 3 $, 3 <j>, same data as holotype, except 
only year date 1936 (2 $, 2 $ in CNC; 1 $, 1 $ in BMNH). 5 $, 3 $, Yugoslavia: Serbia, 
Tuman, 17. v. 1934 (CNC). 

Baranov described this form from both sexes, but without stating the number of specimens. 
A single specimen was designated as 'Typus' in the original publication but without stated 
sex, and Baranov cited the type-locality as 'Dedinje-Bach bei Golubac'; we assume therefore 
that the holotype specimen was from the Dedinje stream, although this name does not 
appear on the holotype data label. 

The CNC collection contains (from Baranov's collection) 5 $ and 3 $ specimens of ander- 
liceki collected at Tuman (Serbia) on 17.V.1934. This locality was cited in the original 
descriptions, though without date, and these specimens are considered to be paratypes. 

0. ornata i. anderliceki is not placed by Rubzov (1959-1964) ; we confirm from holotype 
examination that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 500) gives the publication date of this name as 1936 in error. 

Odagmia ornata (Meigen), form babici Baranov, 1937a : 261 (Croatian), 273 (German). 
Holotype S, Yugoslavia: Macedonia, Skopje, Vardar, 21.1.1927 (CNC). 

Paratypes : 1 <J, 1 $, same data as holotype, except $ without 'Vardar' cited (CNC) . 1 ?, 
same data as holotype, except date 30.x. 1926 (CNC). 1 $, same date as holotype, except 
date 18.1.1927 (CNC); 2 $, same data as holotype, except date 18. xi. 1926 (CNC and BMNH). 
6 (J, 2 ?, same data as holotype, except various dates 1-23.il. 1927 (all in CNC except one $ 
with date 23. ii. 1927 in BMNH). 2 $, same data as holotype, except date 26. v. 1927 (CNC). 
1 ?, same data as holotype, except additional locality Glumovo near Skopje and date 
12.iv.1927 (CNC). 

Baranov described this form from both sexes, but without stating the number of specimens. 
A single specimen, without stated sex, was designated as 'Typus' in the original publication 
and the date given as '21. 1. 1927', no other date being mentioned. We accept all of the above- 
listed paratypes as having this status as they all come from the type-locality or nearby, but 
most of them bear dates not cited by Baranov in the description. Baranov cited the exact 
type-locality as 'Vardar von Skoplje bis zur Miindung der Treska', but this full data does not 
appear on any of the type-specimens. 

O. ornata f. babici is not placed by Rubzov (1959-1964) ; we confirm from holotype examina- 
tion that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 501) gives the publication date of this name as 1936 in error. 

Odagmia ornata (Meigen), subsp. barensis Baranov, 1939a" : 600 (Croatian), 601 (German). 
Syntypes <j>, Yugoslavia: Montenegro, rivulet near Bar (formerly Antivari), viii.1938 (not 
located, presumed lost). 

This subspecies was described from females (number of specimens not stated) reared from 
pupae, but the pupal stage was not itself described. No material of the taxon has been 
found, and the types are presumed lost. 

The name barensis was overlooked by Smart (1945) and omitted from his catalogue of 
world Simuliidae. It is not placed by Rubzov (1959-1964) and remains enigmatic, though 
it may safely be presumed that it correctly applies to some species (probably ornata) of the 
genus-group segregate Odagmia Enderlein). 

Odagmia ornata (Meigen), form bartulici Baranov, 1937a : 262 (Croatian), 274 (German). 
Holotype <$, Yugoslavia: Serbia, Donji Milanovac, 6.V.1936 (CNC). 

Paratypes: 1 <?, 4 $, same data as holotype {$, 3 ? in CNC; 1 $ in BMNH). 1 $, Yugo- 
slavia: Serbia, Golubac, i.vi.1924 (CNC); 1 ?, same locality, date 20.V.1925 (CNC); 1 $, 2 2, 
same locality, date 30.vi.1925 (CNC). 2 $, without data (with Baranov's name labels, 
paratype status presumed). 



BARANOV'S TYPES OF SIMULIIDAE 199 

Baranov described this form from both sexes, but without stating the number of specimens. 
A single specimen, without stated sex, was designated as 'Typus' in the original publication 
and the date given as '6. v. 1936'; the type-locality was given as 'D. Milanovac (der Bach 
iiber welchen der Weg von D. Milanovac nach Greben fiirht)'. Baranov also mentioned the 
locality 'Golubac' but without giving dates for the specimens from this locality (which have 
paratype status). 

O. ornata f. bartulici is not placed by Rubzov (1959-1964); we confirm from holotype 
examination that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 501) gives the publication date of this name as 1936 in error. 

Odagmia ornata (Meigen), form borcici Baranov, 1937a : 261 (Croatian), 273 (German). 
Holotype $ [wings missing], Yugoslavia: Macedonia, Skopje, 17.x. 1926 [publ. as i.xi.1926] 
(CNC). 

Paratypes: 5 $, same data as holotype, except date i.xi.1926 (CNC). 1 cJ, same data as 
holotype, except date 3.vii.i926 (CNC). 1 $, same data as holotype, except date 5.viii.ig26 
(CNC). 1 c?, same data as holotype, except date 11. ix. 1926 (CNC). 1 $, same data as holo- 
type, except date 26.x. 1926 (CNC). 1 cJ, 1 °, same data as holotype, except date 27.x. 1926 
(CNC). 1 °, same data as holotype, except date 30.x. 1926 (CNC). 1 <$, 1 $, same data as 
holotype, except date 4.X1.1926 (BMNH). 2 $, same data as holotype, except date 4.XL1926 
(CNC). 1 ?, same data as holotype, except date 20. xi. 1926 (CNC). 1 S, same data as 
holotype, except date 20. ii. 1927 (CNC). 1 °, same data as holotype, except date 11. iv. 1927 
(CNC). 1 pupal exuvium, same data as holotype, except date 3.viii.ig26 (CNC). 2 °, 
Yugoslavia: Macedonia, Hanrijevo, 27.vi.1926 (CNC). 1 $, Yugoslavia: Macedonia, 
Gradovzi, i.x.1926 (CNC). 

Baranov described this form from both sexes, but without stating the number of specimens. 
A single specimen, without stated sex, was designated as 'Typus' in the original publication 
and the date for this holotype was cited as 'i.xi.1926'. A discrepancy exists in Baranov's 
collection concerning the holotype and its data: the collection contains five specimens on 
two mounts which have the date 'i.xi.1926', but the single female specimen marked as 'Typus' 
by Baranov bears the date '17.x. 1926'; it appears either that Baranov marked the wrong 
specimen as type or that he cited the wrong date in publication. The specimens having the 
date ' i.xi.1926' exist as a pair on one mount and a trio on one mount, and it is unlikely that 
Baranov intended one of these specimens — not clearly separated from the others — to be 
'Typus', and we consider it best to accept the specimen indicated as type by Baranov to be 
the holotype. We hold, therefore, that Baranov inadvertently cited the wrong date in 
publication for this specimen, and that the correct date for the holotype data is '17.x. 1926'. 

The form borcici is one of the 'Vardar-Formenkomplex', i.e. forms of ornata described 
from the geographical area of the Vardar river, Macedonia, and the specimens came from the 
Skopje and Hanrijevo environs according to Baranov's description. The specimens listed 
above, apart from the holotype, are all paratypes, but it should be noted that the CNC 
collection contains in addition two male and two female specimens under the name borcici 
from Baranov's collection that have the data 'Golubaz, io.xi.[i]g24'; as these specimens are 
from Golubac on the Danube in Serbia (outside the area of the 'Vardar-Formenkomplex') 
and this locality is nowhere mentioned in the original description, it is considered that they 
have no type-status. 

O. ornata f. borcici is not placed by Rubzov (1959-1964) ; we confirm from holotype examina- 
tion that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 502, 510) listed borcici with the erroneous spelling borici, and (p. 502) gave 
the publication date as 1936 in error. 

Odagmia ornata (Meigen), form guelminoi Baranov, 1937a : 262 (Croatian), 273 (German). 
Holotype $, Yugoslavia: Serbia, Nis, i.vi.1935 (CNC). 

Paratypes : 3 <?, 2 ?, same data as holotype (CNC, all badly damaged) . 22 <$, 32 ?, Yugo- 
slavia: Serbia, Pukovac, 1935 (CNC, except 2 $, 2 $ in BMNH). 6 $, Yugoslavia: Serbia, 
Tuman, vi.1925 (CNC). 



zoo R. W. CROSSKEY & B. V. PETERSON 

Baranov described this form from both sexes, but without stating the number of specimens. 
The single specimen listed above as holotype was designated as 'Typus' but without stated sex. 

0. ornata f. guelminoi is not placed by Rubzov (1959-1964); we confirm from holotype 
examination that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 505) gives the publication date of this name as 1936 in error. 

Odagmia ornata (Meigen) form nikolici Baranov, 1937a : 262 (Croatian), 274 (German). 
Holotype $, Yugoslavia: Serbia, Nis, A [sicj.iv.1935 (CNC). 

Paratypes: 3 c?. 1 9, same data as holotype (2 $, ? in CNC; 1 <$ in BMNH). 4 <J, 1 $, 
Yugoslavia: Serbia, Kursumlija, 10. iv. 1935 (CNC). 

Baranov described this form from both sexes, but without stating the number of specimens. 
The single specimen cited above as holotype was designated as 'Typus' but without stated 
sex. The three Serbian localities Nis, Pirot, and Kursumlija, were cited in the description, 
but we have found no paratype specimens from the Pirot locality. 

O. ornata f. nikolici is not placed by Rubzov (1959-1964); we confirm from holotype 
examination that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 509) gives the publication date of this name as 1936 in error. 

Odagmia ornata (Meigen), form prima Baranov, 19266 : 184, 189. LECTOTYPE $, by 
present designation, Yugoslavia: Serbia, Tuman, 5.^.1925 (CNC). 

Paralectotypes : 1 (J, same data as lectotype (and on same mount) (CNC). 3 ?, Yugo- 
slavia: Serbia, Golubac, 25.lv. 1925 (CNC); 1 <J, 2 $, same locality, date 26.iv.1925 (CNC); 
1 <J, 1 ?, same locality, date 24. v. 1925 (CNC). 

This form was described from both sexes, pupae and larvae, but we have seen no immature 
stage syntypes and presume that these were lost with the rest of Baranov's alcohol material. 
No localities were cited in the original description, Baranov merely stating that the form was 
present everywhere ('llberall vorhanden') ; all the above-listed specimens can be accepted as 
original syntypes. In addition the CNC collection contains, from Baranov's collection, 
another female specimen from the type-locality (Tuman) but with the collection date '26. iv. 
1927'; as this specimen was collected in the year after form prima was described it is not an 
original syntype. 

O. ornata f. prima is not placed by Rubzov (1 959-1 964) ; we confirm from lectotype examina- 
tion that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

The name is a primary homonym of prima Baranov (form of Odagmia ruficomis), and a 
secondary homonym in Simulium s.l. of prima Baranov (19266, forms of equina L., aurea 
Fries, and stylata Baranov). 

Odagmia ornata (Meigen), form savici Baranov, 1937a : 262 (Croatian), 274 (German)' 
Holotype <j, Yugoslavia: Serbia, Pirot, 12. iv. 1935 (CNC). 

Paratypes: 5 <J, 2 $, same data as holotype (4 3, 1 $ in CNC; 1 <J, 1 $ in BMNH). 

Baranov does not specifically mention characters of the male in the original descriptions, 
but as males with the correct cited date (of which one is labelled by Baranov as 'Typus') 
and from the single cited locality stand with the females in his collection, and as it is known 
that all the other forms of ornata described in the 1937a work were based on both sexes, we 
consider it certain that the males were before Baranov at the time of description and that 
they should be considered to be original syntypes. 

O. ornata f. savici is not placed by Rubzov (1959-1964) ; we confirm from holotype examina- 
tion that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 513) gives the publication date of this name as 1936 in error. 

Odagmia ornata (Meigen), form secunda Baranov, 19266 : 184, 191. Syntypes c?, ?, pupae, 

larvae, Yugoslavia: Serbia, Tuman; & Macedonia, Skopje (not located and presumed lost). 

Baranov described both sexes of this form, and also presumably had the immature stages 

before him at the time of description (as he stated 'Puppe und Larve sind der 0. ornata 



BARANOV'S TYPES OF SIMULIIDAE 201 

prima ahnlich'). No adult syntypes are present in Baranov's collection in CNC, and his 
alcohol material of immature stages was doubtless lost with the rest of his alcohol collection 
at the end of the war in Vienna. So far as we can tell, therefore, all type-material is lost. 

However, it should be noted that Baranov's collection in CNC contains one female specimen 
from Skopje (one of the syntypic localities recorded by Baranov), but this specimen has the 
collection date 'i.xi.1926'; the paper containing the original description of secunda was 
published in March, 1926, and this specimen cannot therefore be an original syntype (it is 
labelled as 'neotypus' by Baranov but was never published as such). The CNC collection 
also contains, from Baranov's collection, labelled in error as cotypes, one female and two male 
specimens named as ornata form secunda and having the data 'Golubac, 26. iv. 1927'; these 
specimens also are not original syntypes as they come from a locality not mentioned in the 
original description (Baranov stated that he had specimens only from Tuman and Skopje) 
and were collected more than a year after the original description had been published. 

0. ornata i. secunda is not placed by Rubzov (1959-1964); in the absence of type-material 
we cannot absolutely confirm that assignment to Odagmia is correct, but this may be presumed 
from the fact that ornata is type-species of Odagmia and Baranov's form secunda is really 
certain to belong to this genus-group concept. 

The name is a primary homonym of secunda Baranov (form of Odagmia ruficornis), and a 
secondary homonym in Simulium s.l. of secunda Baranov (19266, forms of equina L., aurea 
Fries, and stylata Baranov) . 

Odagmia ornata (Meigen), form zagrebiensis Baranov, 1937a : 263 (Croatian), 274 (German). 
LECTOTYPE cJ, by present designation, Yugoslavia: Croatia, Zagreb, 10.ii.1931 (CNC). 

Paralectotypes : 1 S, same data as lectotype, except date 16.1.1931 (CNC). 1 <J. same 
data as lectotype, except date 27.1.1931 (CNC). 1 <J, same data as lectotype, except date 
12.ii.1931 (BMNH). 1 $, same data as lectotype, except date 17.ii.1931 (CNC). 

Baranov described this form from an unstated number of specimens of both sexes and did 
not designate a 'Typus' ; present designation of a lectotype is therefore necessary. In the 
original publication Baranov gave the date range '10-17. ii. 1931' an d this agrees with data 
labels on most of the specimens in his collection ; however there are two specimens with slightly 
different dates (viz. 16. i. and 27.1.1931) from that cited, but we nevertheless regard these as 
original syntypes (they were regarded as types by Baranov and all evidence suggests that 
they were available to him at the time of description). 

O. ornata i. zagrebiensis is not placed by Rubzov (1959-1964); we confirm from lectotype 
examination that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 516) gives the publication date of this name as 1936 in error. 

Odagmia ruficornis Baranov, 19266 : 184, and O. ruficornis Baranov, form prima Baranov, 
19266 : 184, 191. LECTOTYPE $, by present designation, Yugoslavia: Serbia, Golubac, 
27.V.1925 (CNC). 

Paralectotypes: 2 <3, same data as lectotype (CNC). 

Baranov's (19266) description of Odagmia ruficornis and its two forms (prima and secunda) 
is very confused, and some discussion is here necessary. The availability of these names 
rests upon the entry in the key on p. 184 of the original publication and on the descriptive 
matter on p. 191. The key entry reads: 

'26 (29) Fuhler des $ ganz hellrot . . . ruficornis n.sp. 

27 (28) H.-Schenkel hell mit schwarzem Ende . . . ruficornis prima n.f. 

28 (27) H.-Schenkel ganz hell . . . ruficornis secunda n.f.' 
and the 'descriptive' entry reads: 

'14-15. Odagmia ruficornis prima und secunda nn.ff. ??. Ausser durch die roten Fuhler 
unterscheiden sie sich von O. ornata durch eine hellere Korperfarbe und gleichmassige, 
grobere und dichtere, mehr silberige Behaarung. 
Ich habe beide Formen aus Tuman und Golubaz.' 



202 R. W. CROSSKEY & B. V. PETERSON 

The numbers 14 and 15 preceding the descriptive matter are serial numbers in a list of Simuliid 
species (the preceding number 13 referring to Odagmia ornata nitidifrons Edwards and the 
succeeding number 16 referring to Odagmia kondici Baranov). It is evident therefore that 
Baranov was proposing two separate taxa only (No. 14 and No. 15), differing in detail of leg 
colouring but together forming the new species ruficornis distinguished by its red antennae. 
From this it is plain that form prima was intended to be the typical or nominate form, and 
therefore that the species ruficornis is based upon the same type-material as form prima. 
Baranov's collection in CNC contains no specimens labelled as prima but does contain speci- 
mens labelled simply as ruficornis, and these specimens if accepted as syntypes are auto- 
matically types of both ruficornis and of ruficornis f. prima. 

Unfortunately the status of the existing specimens in Baranov's collection is not absolutely 
certain, because all of them are males and Baranov's very deficient descriptive matter and 
key appear to mention only the female. It is well known, however, that Baranov must have 
had on many occasions specimens in front of him that he failed to note in publication, and 
that many types of discrepancy exist between cited information in his publications and the 
sex and data of specimens. In the case of ruficornis and ruficornis f. prima we here take the 
view that the three $ specimens in Baranov's collection and labelled by him as types are 
acceptable as being original syntypes, and we have designated a lectotype from them 
accordingly. 

The name ruficornis Baranov is a junior secondary homonym in Simulium s.l. of Simidium 
ruficorne Macquart, 1838, and Smart (1944 : 133) published the replacement name baracorne 
Smart for the preoccupied ruficornis Baranov. Rubzov (1963 : 472-474) places ruficornis 
Baranov, under the name Odagmia baracornis (Smart), as a valid species of the ornata-group 
of Odagmia; we confirm from lectotype examination that it is correctly assignable to the 
genus-group segregate Odagmia Enderlein. 

Odagmia ruficornis Baranov, form secunda Baranov, 19266 : 184. LECTOTYPE $, by 
present designation, Yugoslavia: Serbia, Golubac, 27.V.1925 (CNC). 

Paralectotype : 1 $, Yugoslavia: Serbia, Negotin, ix.1924 (CNC). 

Baranov (19266 : 191) only mentioned the localities Tuman and Golubac with reference to 
ruficornis forms prima and secunda but we think it reasonable to accept the specimen from 
Negotin (listed above as paralectotype) as an original syntype; Negotin lies in the same 
general area of north-east Serbia as the cited localities, and the specimen has Baranov's 
label as a type; we infer that the specimen was available to Baranov at the time of description. 

O. ruficornis f. secunda is cited by Rubzov (1963 : 473) in the section headed 'Variabilitat' 
under the name Odagmia baracornis baracornis Smart, and he evidently does not recognize 
separate status from the nominal taxon, prima ( = ruficornis Baranov s.str., = baracornis 
Smart). We confirm from lectotype examination that secunda is correctly assignable to the 
genus-group segregate Odagmia Enderlein. 

The name is a primary homonym of secunda Baranov (form of Odagmia ornata), and a 
secondary homonym in Simulium s.l. of secunda Baranov (19266, forms of equina L., aurea 
Fries, and stylata Baranov). 

Odagmia tenuitarsus Baranov, 1937a : 264 (Croatian), 276 (German). Holotype ?, Yugo- 
slavia: Croatia, Zagreb, 9.^.1929 (CNC). 

This species is not placed by Rubzov (1 959-1 964). We confirm from holotype examination 
that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

The name is a junior secondary homonym in Simulium s.l. of Simulium tenuitarsus Puri, 
!933» an d Smart (1945 : 528) has published the replacement name baranovi Smart for the 
preoccupied tenuitarsus Baranov. 

Smart (1945 : 514) gives the publication date of tenuitarsus Baranov as 1936 in error. 

Simulium agnatum Baranov, 1937a : 259 (Croatian), 272 (German). Holotype ?, Yugo- 
slavia: Serbia (CNC). 



BARANOVS TYPES OF SIMULIIDAE 203 

Paratype: 1 $, Yugoslavia: Bosnia, Sarajevo, 1932 (CNC). 

Baranov described this species only from the two female specimens cited above, which he 
designated as 'Holotypus' and 'Paratypus' respectively. 

This species is not placed by Rubzov (1959-1964); we confirm from holotype examination 
that it is correctly assignable to Simulium Latreille s.str. 

Smart (1945 : 500) gives the publication date of this name as 1936 in error. 

Simulium begbunaricum Baranov, 1924 : 65. Syntypes <$ [? also ?], Yugoslavia: Serbia, 
Golubac & Kucevo & Zajecar (lost). 

The type-material of this species was preserved in alcohol and was lost in Vienna at the 
end of the war (Baranov i.litt. to Shewell). Although the type-material is lost the identity 
of begbunaricum is known from the footnote in Baranov's (19266 : 183) paper in which he 
stated that 'Die Art 5. begbunaricum Bar. 1924 war eine Mischart und zerfiel in equina- 
Formen', a statement that we take clearly to imply that begbunaricum is a synonym of 
Simulium equinum (Linnaeus) ; this in turn confirms that the name applies to a species of the 
genus-group segregate Wilhelmia Enderlein, to which equinum belongs. The name is not 
given by Rubzov (1959-1964). 

The original description of begbunaricum was published in Serbian by Baranov (1924) in 
the Yugoslav journal Glasnik Ministarstva Poljoprivrede i Voda (Vol. 2, No. 7, p. 65), but in 
the following year Baranov (1925) again described the species as new, citing it as 'n.sp.' 
in both a key in Serbian (pp. 6 & 7 of the 1925 paper) and in a German description (p. 10 of the 
1925 paper). Smart (1945 : 501) overlooked the 1924 paper and cited the German descrip- 
tion on p. 10 of the 1925 paper as the original description in error; Smart (loc. cit.) also in- 
advertently mis-spelt the name as begbungaricum . (This curious specific name alludes to the 
cliff-cave of 'Beg-Bunar' on the Danube, from which according to local superstition the 
infamous Golubatz-fly is supposed to emerge, and begbunaricum is the correct spelling.) 

Simulium brnizense Baranov, 1924 : 66. LECTOTYPK ., by present designation, Yugo- 
slavia: Serbia, near Golubac, Brniza [on label 'Brnjica'], 1924 (CNC). 

Paralectotypes : none located, presumed lost. 

The original Serbian description of this species appears to be based mainly on the male and 
includes a figure of the $ hypopygium, but Baranov makes it clear (especially in the second, 
German, description in 1925 mentioned below) that he had several specimens reared from 
pupae and these doubtless included females. We think it justified, therefore, to consider the 
one existing specimen in Baranov's collection in CNC that has the appropriate data (Brniza, 
1924) and bears Baranov's label as 'Typus' as being an original syntype; we here designate it 
as lectotype. No male syntypes exist in Baranov's collection and these must be presumed 
lost. 

The CNC collection contains specimens from Baranov's collection standing under the name 
brnizense together with the lectotype, but these additional specimens have no type-status, 
even though labelled as cotypes by Baranov (as none are from the type-locality and all were 
collected after the date of publication) : they comprise four males and two females from Mace- 
donia with the following data: 1 <J, Treska, 9.XL1926; 2 <3, Vardar, 10. xi. 1926; 1 ?, Skopje, 
11. xi. 1926; 1 (J, Skopje, 17. xi. 1926; 1 $, Kabajep [spelling partly illegible, uncertain], 20. v. 1927. 

The original description of brnizense was published in Serbian by Baranov (1924) in the 
Yugoslav journal Glasnik Ministarstva Poljoprivrede i Voda (Vol. 2, pt. 7, p. 66), but in the 
following year Baranov (1925) again described the species as new, citing it as 'n.sp.' in both a 
key in Serbian (pp. 6 & 7 of the 1925 paper) and in a German description (p. 10 of the 1925 
paper). Smart (1945 : 502) made a very confused entry for the species in his world cata- 
logue of Simuliidae, giving two different spellings and two different references for the same 
species: his first reference to 'brizensis [sic] Baranov (1924 : 66)' is in error for the spelling of 
the name but is correct for the bibliographic reference, but his second reference to 'bruigense 
Baranov (1925 : 10)' is wrong for both spelling of the name and the reference. The reference 



2o 4 R. W. CROSSKEY & B. V. PETERSON 

given by Smart (loc. cit.) against the mis-spelling bruigense refers to Baranov's German 
description of bmizense in the later (1925) paper and not to the first (original) Serbian des- 
cription. 

We confirm from lectotype examination that bmizense is correctly assignable to the genus- 
group segregate Wilhelmia Enderlein, where Baranov (19266 : 187, and subsequent publica- 
tions) himself placed it. Rubzov (1962 : 401), following earlier authors and using the mis- 
spelling brizensis, places the name as a synonym of falcula Enderlein (a supposed subspecies 
of equina (L.)). There are several references in the literature to Baranov's name with the 
spelling brizensis, but the name alludes to Brniza near Golubac in Serbia and Baranov's 
original spelling is correct. 

Simulium columbaczense (Schoenbauer), race litorale Baranov, 19376 : 159, 164. LECTO- 
TYPE cJ, by present designation, Yugoslavia: Serbia, Golubac, 26. v. 1936 (CNC). 

Paralectotypes : 2 ?, same data as lectotype, except date 22.V.1928 (CNC). 1 <$, 20 §, 
same data as lectotype, except date 25. v. 1928 (CNC). 9 <J, 62 $, same data as lectotype 
(7 cJ, 60 ?, in CNC; 2 <J, 2 ? in BMNH). 

Baranov did not publish a formal description of this race, but the name is nomenclatorially 
available from its first publication by Baranov (19376) because characteristics are cited of the 
adult which differentiate litorale; all adult specimens cited above are acceptable as original 
syntypes and a lectotype is designated from them. Race litorale was named for those Simu- 
lium columbaczense (Schoenbauer) that breed in shallow waters of the Danube near Golubac, 
and in a later paper Baranov (1939a : 122) was very precise about the exact type-locality, 
stating that 'die typische Lokalitat dieser Rasse [i.e. litorale] ist die Insel in der Nahe der 
Dampferanlegestelle bei Golubac' ; this full detail is not however indicated on the type- 
material. Baranov referred to litorale as an ecological race ('okologische Rasse') morpho- 
logically distinct from another ecological race (profundale) with which it was allopatric, since 
the profundale race occurred in the depths of the Danube at Donji Milanovac; as Baranov 
did not recognize subspecies of columbaczense, and as he referred litorale and profundale to be 
characteristic of particular geographical areas, the names of these ecological races are not 
interpreted as infrasubspecific in the terms of Article 45 (d) of the International Code of 
Zoological Nomenclature, 1961; they are therefore available species-group names with status 
under the Code. 

No authors except Baranov have seen the need to recognize named 'races' of the Golubatz 
fly, and we concur in this view. The name litorale is a synonym of Simulium colombaschense 
(Fabricius) (syn. 5. columbaczense (Schoenbauer)), the type-species of Simulium Latreille, 
without separate species-group status. Rubzov (1959-1964) omits the name litorale. 

Simulium columbaczense (Schoenbauer), race profundale Baranov, 19376 : 159, 164. 
LECTOTYPE $, by present designation, Yugoslavia: Serbia, Donji Milanovac, 9.V.1936 

(CNC). 

Paralectotypes: 23 $, 26 $, same data as lectotype (21 $, 24 <j> in CNC; 2 $, 2 $ in BMNH). 

Baranov did not publish a formal description of this race, but the name is nomenclatorially 
available from its first publication by Baranov (19376) because characteristics are cited of the 
adult which differentiate profundale; all adult specimens cited above are acceptable as 
original syntypes and a lectotype is designated from them. Race profundale was named for 
those Simulium columbaczense (Schoenbauer) that breed in the depths of the Danube near 
Donji Milanovac, and in a later paper Baranov (1939a : 122) was very precise about the exact 
type-locality, stating that 'die typische Lokalitat ist bei Donji Milanovac, hinter den 
Traversen der Flussregulierung in einer Tiefe von 4-6 m, doch gibt es auch Fundorte von 
einer Tiefe die 20 m uberschreitet' ; this full data does not of course appear on the data on the 
adult syntypes, which are simply labelled as from D. Milanovac. 

Baranov referred to profundale as an ecological race ('okologische Rasse') morphologically 
distinct from another ecological race {litorale) with which it was allopatric, since the litorale 



BARANOV'S TYPES OF SIMULIIDAE 



205 



race occurred in the Danube in shallow waters near Golubac; the name pvofundale has status 
in nomenclature for the reasons outlined under litorale (see above). 

No authors except Baranov have seen the need to recognize named races of the Golubatz 
fly, and we concur in this view. The name profundale is a synonym of Simulium colom- 
baschense (Fabricius) (syn. S. colnmbaczense (Schoenbauer)), the type-species of Simulium 
Latreille, without separate species-group status. Rubzov (1959-1964) omits the name 
profundale. 

Simulium djerdapense Baranov, 1937a : 258 (Croatian), 270 (German). Holotype $, 
Yugoslavia: Serbia, Golubac, 28. iv. 1936 (CNC). 

Baranov did not state the number of specimens in the original description, which was 
based solely on the female, and in the absence of any evidence to the contrary we accept the 
single $ specimen in Baranov's collection in CNC as holotype; its data agree with that cited 
in the original publication. 

This species is not placed by Rubzov (1959-1964); we confirm from holotype examination 
that it is correctly assignable to the genus-group segregate Odagmia Enderlein. 

Smart (1945 : 504) gives the publication date of this name as 1936 in error. 

Simulium reptans (Linnaeus), form calopum Baranov, 19266 : 184, 189. LECTOTYPE ?, 
by present designation, Yugoslavia: Serbia, [River] Timok, ix.1924 (CNC). 

Paralectotype : 1 $, same data as lectotype (CNC). 

In the case of this nominal taxon there is serious discrepancy between the specimens that 

are labelled as types of calopum by Baranov in his collection in CNC and the information 

published in the original description, and it is necessary to annotate this in some detail. The 

entire description of Simulium reptans calopum n.f. (in Baranov, 19266 : 189) reads as follows: 

'Diese Form unterscheidet sich von beiden vorhergehenden in beiden Geschlechtern 

durch die helleren Beine. Hypopygium des Mannchens (Fig. 5) mit fiir reptans charak- 

teristischem Griffel und starker Biirste. Ich habe sie aus dem Timok und dem Vardar 

und aus einem grosseren Bach neben Svilajnaz (Serbien).' 

From this description two things are certain, firstly that the description was based on both 
sexes and secondly that the original (type) material was from the rivers Timok (in Serbia) 
and Vardar (in Macedonia), and from a stream at Svilajnaz ( = Svilajnac) in Serbia. In all, 
Baranov's collection in CNC contains five (J, seven $ and three pupal (exuviae) specimens, 
each bearing Baranov's cotype label, standing under calopum, but of these only two females 
are true original syntypes: these two specimens are from the river Timok, one of the cited 
type-localities, and have the collection date 'IX. 1924', early enough to pre-date the original 
description. 

The remaining specimens of calopum in the Baranov collection have no type-status, either 
because they are not from an original type-locality or because the collecting dates post-date 
the time of description (or because both date and locality conflict). Apart from the two 
Timok syntypes (here designated as lectotype and paralectotype) the data of the other 
specimens that lack type-status are as follows: Serbia, Golubac; 1 <J, 1 $, date a. [sic] vi.1924; 
1 c?, date 22. iv. 1925; 1 ?, date 13. v. 1925; 1 $, date 25.lv. 1927. Serbia, Brniza; 1 $, date 
30.iv.1927. Serbia, Tuman; 1^,1$, date 26. iv. 1927. Macedonia, Skopje; 1 9, date 
11. iv. 1927 (labelled as type by Baranov) ; 1 (J, date 11. ix. 1927; 3 pupal exuviae, date 9.^.1927. 
The last-named locality, Skopje, is on the river Vardar (one of the type-localities cited by 
Baranov) and on the basis of locality alone the specimens from Skopje appear at first to be 
syntypes, but as the original description was published in 1926 (March) and the specimens 
were not collected until 1927 they cannot be original material; likewise the specimens from 
Golubac, Brniza and Tuman are not original syntypes as they are not from the type-localities 
and most of them also were collected after the time of description. 

5. reptans i. calopum is not placed by Rubzov (1959-1964); we confirm from lectotype 
examination that it is correctly assignable to Simulium Latreille s.str. 



2 o6 R. W. CROSSKEY & B. V. PETERSON 

Simulium reptans (Linnaeus), subsp. glumovoense Baranov, 1937a : 257 (Croatian), 270 
(German). LECTOTYPE $, by present designation, Yugoslavia: Macedonia, Skopje, 
Glumovo, 12. iv. 1927 (CNC). 

Paralectotypes : 2 ?, same data as lectotype (CNC). 1 $ ex pupa (cocoon only remaining), 
Yugoslavia: Macedonia, [River] Treska, Glumovo, 6.^.1927 (pupa), 10. iv. 1927 (emerged 
adult) (CNC). 

This subspecies was described only from the female. The number of specimens was not 
clearly indicated, but Baranov cited the dates 6.iv. and 10. iv. 1927 near the beginning of the 
Croatian description (p. 257) and 12. iv. 1927 at the end, implying more than one specimen 
(although in the abbreviated German description on p. 270 he mentioned only the date 
1 2. iv. 1927). There are four specimens in Baranov's collection in CNC conforming to the sex 
and various cited dates and these are clearly all original syntypes and are listed above; in 
addition there is one male specimen from the type-locality (Skopje, Glumovo) but as the male 
was not described and the specimen has a conflicting date (13.lv. 1927) it is not accepted as 
having any type-status. 

This subspecies is not mentioned by Rubzov (1959-1964); we confirm from lectotype 
examination that it is correctly assignable to Simulium Latreille s.str. 

Smart (1945 : 505) gives the publication date of this name as 1936 in error. 

Simulium reptans (Linnaeus), form ornatoide Baranov, 19266 : 184, 189. Holotype or 
syntypes $, Yugoslavia: Serbia, Tuman, 12.vi.1925 (lost). 

The type-material of this species was preserved in alcohol and was lost in Vienna at the 
end of the war (Baranov i.litt. to Shewell). 

5. reptans f . ornatoide is not placed by Rubzov (1959-1964) . In the absence of type-material 
the name is enigmatic, but presumably applied without doubt to a species of Simulium 
s.str., in which genus-group segregate S. reptans (Linnaeus) belongs. 

Simulium reptans (Linnaeus), subsp. pseudocolumbaczense Baranov, 1937a : 255 
(Croatian), 269 (German). Holotype $, Yugoslavia: Serbia, Golubac, Donau [ = R. Danube], 
i.v.1936 (CNC). 

This subspecies was described from a single specimen (the above-listed holotype) reared 
from the pupa. Baranov makes it clear that he had only one specimen from statements in 
both Croatian and German descriptions. 

This subspecies is not mentioned by Rubzov (1959-1964); we confirm from holotype 
examination that it is correctly assignable to Simulium Latreille s.str. 

Smart (1945 : 512) gives the publication date of this name as 1936 in error. 

Simulium reptans (Linnaeus), subsp. pukovacense Baranov, 1937a : 256 (Croatian), 269 
(German). LECTOTYPE $, by present designation, Yugoslavia: Macedonia, Pukovac, 
23V.I935 (CNC). 

Paralectotypes: 1 6*. 5 ?> same data as lectotype (<J, 4 $ in CNC; 1 $ in BMNH). 

This subspecies is not mentioned by Rubzov (1 959-1 964); we confirm from lectotype 
examination that it is correctly assignable to Simulium s.str. 

Smart (1945 : 512) gives the publication date of this name as 1936 in error. 

Simulium reptans (Linnaeus), subsp. tumanicum Baranov, 1937a : 255 (Croatian), 269 
(German). Holotype ?, Yugoslavia: Serbia, Tuman, 16.iv.1926 (CNC). 

In the original description Baranov cited one female specimen from Tuman (locality to 
which the name refers) with date 16. iv. 1926, and the specimen with this data in the Baranov 
collection is certainly the holotype. However it should be noted that CNC collection con- 
tains three other specimens of tumanicum from Baranov's collection which have no type- 
status, viz. 2 o\ Tuman, 9.^.1926, and 1 $, Serbia, Golubac, 13.lv. 1926; there is no evidence 
that Baranov had the male at the time of description, these dates were not cited, and no 



BARANOV'S TYPES OF SIMULIIDAE 207 

specimens were mentioned from Golubac, and we conclude that the specimens lack type- 
status even though labelled as cotypes by Baranov. 

This subspecies is not mentioned by Rubzov (1959-1964); we confirm from holotype 
examination that it is correctly assignable to Simulium Latreille s.str. 

Smart (1945 : 515) gives the publication date of this name as 1936 in error. 

Simulium reptans (Linnaeus), subsp. vardaricum Baranov, 1937a : 256 (Croatian), 270 
(German). Holotype ?, Yugoslavia: Macedonia, Skopje, 11.vi.1926 (CNC). 

Paratypes: 1 pupal exuvium, same data as holotype, except date 31.vii.1926 (CNC). 1 
cocoon, same data as holotype, except date 9.^.1927 (CNC). 

In the original Croatian description Baranov briefly described the larva and pupa of this 
subspecies from the Vardar River near Skopje, and based the description of the adult female 
on one specimen reared from the pupa which emerged on n.vi.1926 and was cited as type. 
As the pupa of this subspecies was described, as well as the female, we accept the pupal pelt 
and the empty cocoon standing with the holotype in Baranov 's collection as paratypes (see 
above) ; they came from the type-locality, but Baranov did not mention dates for his pupal 
material in the description. 

This subspecies is not mentioned by Rubzov (1959-1964); we confirm from holotype 
examination that it is correctly assignable to Simulium Latreille s.str. 

Smart (1945 : 515) gives the publication date of this name as 1936 in error. 

Simulium (Nevermannia) serbicum Baranov, 1925 : 6, 7 (Serbian), 9 (German). Syntypes 
o", $, pupae, Yugoslavia: East Serbia, stream near Yrashegranaz, 21. ix. 1924 (not located, 
probably all lost). 

Baranov described this species from one female and three male specimens which had been 
reared from pupae. The pupae were stated to recall those of small ornatum, but to differ 
by having only four gill filaments. No type-specimen was designated, and the four reared 
adults plus pupae therefore had syntype status. 

In the description of the male Baranov recorded Teh besitze nur Alkohol Material'; the 
alcohol material from Baranov's collection is known to be lost and there is no extant male or 
pupal syntype material. The one female syntype was not stated to be in alcohol and may 
have been a pinned specimen, but it, too, appears to be lost. It must be noted, however, that 
Baranov's list of specimens sold to the Department of Agriculture, Ottawa, in 1954 mentioned 
ones specimen of serbicum, and that this specimen was marked off as present in the collection 
when it was received at Ottawa; the sex was not recorded. Whether this specimen was the 
single female syntype cannot be ascertained, as the specimen cannot now be found in the 
Baranov collection in Ottawa; it appears to have been lost, assuming it was in fact received 
in Ottawa, as records show that no specimen of serbicum was sent out on loan from the collec- 
tion (failure to return a loan specimen therefore does not account for absence of the serbicum 
specimen from Baranov's collection). 

Sherban (1961 : 677) reported that he had studied Baranov's 'holotype' of serbicum and 
found that it did not belong to the same species as serbicum Baranov in the sense of Rubzov. 
We are puzzled by this statement, as we do not think it possible that the specimen to which 
Sherban referred can possibly be an original type-specimen: most of the original material 
was in alcohol and is known to be lost, there are no Baranov simuliid types existing in Yugo- 
slavia or elsewhere in south-east Europe so far as we know, and Sherban did not see the one 
specimen of serbicum in Ottawa from Baranov's collection (see above paragraph). We have 
not been able to obtain information from Dr Sherban but we tentatively conclude that the 
specimen he saw was not one of Baranov's original serbicum specimens. It appears instead 
to have been a specimen from the river Medoviza in Yugoslavia which Rubzov in error con- 
sidered to be a serbicum type, because Rubzov (1962 : 380) in his account of serbicum makes 
the statement 'Typus aus dem Fluss Medoviza (Jugoslawien)'; Rubzov's statement of the 
type-locality is completely at variance with anything indicated by Baranov, and his 'Typus' 
cannot in reality, whatever the specimen may be, have any type-status. Sherban mentions 



208 R. W. CROSSKEY & B. V. PETERSON 

the Medoviza locality and Rubzov's specimen and appears to have confused this with Bara- 
nov's 'type'. That Sherban was following Rubzov's interpretation of the 'type' of serbicum 
seems evident from the fact that he repeated Rubzov's (1956 : 521) error of giving Pseu- 
donevermannia as the original genus : serbicum was actually described by Baranov in Simnlium 
(Nevermannia), and the description in fact predated the erection of the segregate Pseu- 
donevermannia Baranov, 1926. 

Shortly after the description Baranov (19266, 1927a and 1929) placed serbicum in Never- 
mannia, regarding the latter as a genus, and aggregated it with aareum Fries. Nevermannia 
is now regarded as a synonym of Eusimulium, to which segregate serbicum (though the type- 
material is lost) undoubtedly belongs: Baranov's (1925) figure of the male hypopygium and 
his figure of the branching of the 4-filamented pupal gill (Baranov, 1927a) confirm that 
the species belongs in the aureum-group of Eusimulium, where Rubzov (1956 : 521; 1962 : 
380) places it. 

Smart (1945 : 513) cited Nevermannia as the original genus for serbicum, whereas the 
original combination was Simulium (Nevermannia) serbicum. Smart (op. cit.), following 
Enderlein, gave serbicum as a synonym of kerteszi Enderlein, but this synonymy is not upheld 
by Rubzov and is almost certainly wrong (the two nominal species correctly belonging in 
different species-groups) . 

Zivkovic (1966 : 263), in her account of the aureum-group in Yugoslavia, mentions that 
Sherban considered serbicum in the sense of Rubzov to be a misidentification of the true 
serbicum Baranov, and uses the name Simulium rubzovianum Sherban (which Sherban, 1961, 
proposed for serbicum of Rubzov, not Baranov) for a valid species of the group; she does not, 
however, provide any information on the identity of the true serbicum Baranov (though 
almost certainly this name must apply to, and have priority for, one of the Yugoslav species 
of the aureum-group to which she refers). 

Wilhelmia equina (Linnaeus), form prima Baranov, 19266 : 184, 185. LECTOTYPE J, 
by present designation, Yugoslavia: Serbia, Tuman, lower Danube funtere Donau'j, 
23. vi. 1924 (CNC, with pupal exuvium). 

Paralectotypes : 1 $, Yugoslavia: Serbia ('Ost-Serbia'), Tuman, 12. vi. 1925 (CNC). 3 
o\ 4 $, Yugoslavia: Serbia, Tuman, 9.VL1925 (CNC). 

There is some confusion in Baranov's collection concerning the true type-material of this 
nominal taxon, for the collection contains specimens that cannot be types although labelled 
as such by Baranov and at the same time contains specimens that we believe to be types 
though not labelled as such. Detailed annotation is therefore necessary. 

Form prima of equina is a nominal species-group taxon of which the syntypes are the original 
male, female, pupal and larval specimens very briefly described by Baranov from specimens 
obtained at Tuman and Treska; the only information in the original description 
concerning locality and date is contained in Baranov's statement Teh sammelte 
sie massenhaft im Juni in Tuman (Serbien) und im August-September in 
Treska (Mazedonien)'. In practice Baranov used the name prima for what would now be 
referred to as the typical or nominate form, so that his equina prima is synonymous with the 
entity we would now call equina equina. His collection in CNC contains specimens standing 
under the name Wilhelmia equina and others named as equina form prima, but we believe that 
some of the true original syntypes of prima are contained among the specimens named simply 
as equina (not surprising in view of the fact that Baranov's typical equina concept was the same 
as prima), and that his specimens named as types of prima can have no such status. 

The specimens named and labelled as types of prima were all collected after the date of 
publication of the name (March, 1926), and therefore cannot be original syntypes in spite of 
Baranov's labels; furthermore they were all collected at Skopje, which is not one of the cited 
original localities. In all they are comprised as follows: 1 o\ 1 $, Skopje, Vardar River, 
7.XL1926; 1 $, Skopje, 8.ix.i926; 1 $, Skopje, 4.XL1926. We consider all of these specimens 
to be later collected material erroneously labelled as types. 



BARANOV'S TYPES OF SIMULIIDAE 209 

Among the material named simply as equina, however, we have found nine specimens from 
Tuman (one of the originally cited type-localities) that bear collecting dates June, 1924, and 
June, 1925, and we believe that these specimens (which must have been available to Baranov 
at the time of description of prima) represent part of the original prima type-material; their 
data fits with Baranov's statement 'im Juni in Tuman' and they were collected well before 
the publication date of the name. Accordingly we accept these nine specimens as original 
syntypes and designate a lectotype from them (see above, where full data of the lectotype 
and remaining syntypes, i.e. paralectotypes, are indicated). 

The equina material in Baranov's collection contains nine female specimens from Treska 
(the other type-locality that Baranov cited), but as all have data post-dating publication 
none of them are syntypes of prima. We have not located any adult syntypes from this 
locality, and these we believe lost; Baranov's larval and pupal material is also presumed lost 
except for the pupal exuvium of the lectotype and of two of the paralectotypes. 

W. equina f. prima is not mentioned by Rubzov (1959-1964). The name was clearly used 
by Baranov for the typical or nominate equina and is therefore to be treated as a synonym of 
equina Linnaeus, a species-group taxon of the genus-group segregate Wilhelmia Enderlein. 
This is here confirmed from examination of the lectotype. The name is a primary homonym 
of prima Baranov (form of Wilhelmia stylata), and a secondary homonym in Simulium s.l. 
of prima Baranov (19266, forms of aurea Fries, ornata Meigen and ntficomis Baranov). 

Wilhelmia equina (Linnaeus), form quarta Baranov, 19266 : 184. LECTOTYPE cJ, by 

present designation, Yugoslavia: Serbia, Golubac, e [sic] vi.1924 (CNC). 

Paralectotype : 1 o\ same data as lectotype (but label with capital 'E') (CNC). 

Smart (1945 : 512) cites p. 186 in Baranov (19266) for the original description of this form 
but differentiating features of quarta are not clearly stated on this page, and the availability 
of the name rests upon Baranov's entry in the key on p. 184, where the form is differentiated 
by its pale legs. Baranov did not indicate the sex and presumably his statement of 'Beine 
hell' is meant to apply to both sexes; he certainly also had larvae and pupae at the time of 
description as the immature stages of forms quarta, secunda and tertia are referred to on 
p. 186. 

The only existing specimens that can be considered original syntypes are the two males 
recorded above as lectotype and paralectotype. Baranov did not cite any localities for 
quarta but on p. 184, before the key, he mentions that the paper as a whole is based on speci- 
mens from north-east Serbia and from the surroundings of Skopje; the specimens from Golubac 
(north-eastern Serbia) were collected in June, 1924, and can therefore be considered original 
syntypes because of agreement of both locality and date with the publication. But the 
Baranov collection in CNC also contains two $ specimens from Skopje, with dates 3.XL1926 
and 4.xi.ig26 respectively, which have no type-status (although they come from a cited type- 
locality) ; the original description was published in March, 1926, and these females collected 
in November, 1926, cannot therefore be syntypes. 

W. equina f. quarta is not placed by Rubzov (1959-1964); we confirm from lectotype 
examination that it is correctly assignable to the genus-group segregate Wilhelmia Enderlein. 

Wilhelmia equina (Linnaeus), form secunda Baranov, 19266 : 184. Syntypes $, ?, pupae, 
larvae, Yugoslavia (locality and dates uncertain) (lost). 

The type-material of this form was preserved in alcohol and was lost in Vienna at the end 
of the war (Baranov i.litt. to Shewell). It is clear from Baranov's statements on p. 186 
of the original work that he knew the larval and pupal stages as well as the adults differen- 

i tiated in the key on p. 184 (the availability of the name rests on this entry, not on p. 186 
cited for the original description by Smart (1945 : 513)). 

Rubzov (1962 : 412) places Wilhelmia secunda Baranov as a valid species, says that the 
aquatic stages are unknown ('Wasserbewohnende Stadien unbekannt') and says that the 
principal form ('Stammform') is from the Danube. In the absence of any statement of 
locality for W. equina i. secunda and -with the loss of the type-material there is no evidence 



210 R. W. CROSSKEY & B. V. PETERSON 

that firmly substantiates Rubzov's placement and statements; certainly the early stages were 
known to Baranov at the time of description, as on p. 186 of the work containing the descrip- 
tion he wrote (under the heading '2-4. Wilhelmia equina secunda, tertia und quarta ff.nn.') 
'Sie haben ^>nma-ahnliche Larven. Die Puppen sind mit Besonderheiten in Tubenbau. 
Alle drei Formen haben kiirzere und dickere Tuben als prima' . 

The name secunda is a primary homonym of prima Baranov (form of Wilhelmia stylata), 
and a secondary homonym in Simulium s.l. of secunda Baranov (19266, forms of aurea Fries, 
ornata Meigen and ruficornis Baranov). All five uses of the name secunda were published 
be Baranov (19266) in the same work; as one of these, viz. secunda as form of equina, has been 
ayopted as the valid name of a supposed species by Rubzov (1956 : 570; 1962 : 412) we 
adcept this use of the name as determining the senior homonym. 

Wilhelmia equina (Linnaeus), form tertia Baranov, 19266 : 184. Syntypes £, ? ?, pupae, 
larvae, Yugoslavia (locality and date uncertain) (lost) . 

The type-material of this form was preserved in alcohol and was lost in Vienna at the end 
of the war (Baranov i.litt. to Shewell). It is clear from Baranov 's statements on p. 186 of 
the original work that he knew the larval and pupal stages as well as the adult male (the 
male is differentiated in the key on p. 184 and the availability of the name rests on this entry, 
not on p. 186 cited for the original description by Smart (1945 : 514)); there is no mentior 
of the female, and it is not certain whether there were females in the type-material. There 
is no indication in the description of the type-locality or dates of the original material. 

Rubzov (1962 : 401) places Wilhelmia WmBaranovasa valid species, and states {op. cit.: 
402) 'Typus aus der Donau, von uns nicht gesehen'. We know of no evidence, and there is 
none in the original description, that the Danube was the type-locality though this is cer- 
tainly possible, for the river borders north-eastern Serbia which was one of the areas mentioned 
rather vaguely by Baranov (19266) in his introductory remarks. According to Rubzov the 
aquatic stages are unknown, but they were certainly known to Baranov at the time of descrip- 
tion, for under the heading (p. 186) '2-4. Wilhelmia equina secunda, tertia und quarta ff.nn.' 
he wrote 'Sie haben ^n'ma-ahnliche Larven. Die Puppen sind mit Besonderheiten in Tuben- 
bau. Alle drei Formen haben kiirzere und dickere Tuben als prima' . 

Wilhelmia stylata Baranov, 19266 : 184, and W. stylata Baranov, form prima Baranov 
19266 : 184, 186. Syntypes $, Yugoslavia: Macedonia, Treska (lost). 
The situation with these names and with W . equina f. secunda exactly resembles that with 
Odagmia ruficornis Baranov and its forms. Baranov's (19266) description of stylata and its 
two forms is confused, and some discussion is necessary. The availability of these names 
rests upon the entry in the key on p. 184 of the original publication and on the descriptive 
matter on p. 186. The key entry reads: 

'7 (10) Adminiculum mit Griffel. Biirste unfrei . . . stylata n.sp. 

8 (9) Beine fast ganz dunkel . . . stylata prima n.f. 

9 (8) Beine teilweise hell . . . stylata se[c\unda n.f.' 
and the 'descriptive' entry reads : 

'5-6. Wilhelmia stylata prima und secunda ff.nn. Ich habe nur wenige Exemplare, 
welche ich aus in Treska gesammeltem Matarial zuchtete. Ich kenne nur Mannchen, 
die man nur mit Hilfe der Hypopygiumpraparation v. equina unterscheiden kann 
(Fig. 2)'. 
The numbers 5 and 6 preceding the descriptive matter are serial numbers in a list of Simuliid 
species (the preceding numbers 2-4 referring to Wilhelmia equina forms secunda, tertia and 
quarta, and the succeeding number 7 referring to Wilhelmia brnizensis) . It is evident there- 
fore that Baranov was proposing two separate taxa only (No. 5 and No. 6) differing only in 
detail of leg colouring as indicated in the key but together forming the new species stylata 
distinguished by its adminiculum. From this it is plain that form prima was intended to be 
the typical or nominate form, and therefore that stylata and prima are nomenclaturally one 
and the same taxon, having the same type-material. In fact the type-material of this taxon, 



BARANOV'S TYPES OF SIMULIIDAE 211 

stylata = prima, was preserved in alcohol (except for the genitalia preparation mentioned by 
Baranov) and was lost in Vienna at the end of the war (Baranov i.litt. to Shewell). The 
genitalia slide (s) are also presumably lost. 

Smart (1945) cites p. 185 for the original description of stylata and its forms; this is in error 
and details of these taxa appear on pp. 184 and 186. 

Rubzov (1962 : 407) places Wilhelmia stylata Baranov as a synonym of Wilhelmia lineata 
(Meigen). This is accepted as correct in the absence of any evidence to the contrary. 

The name prima is a primary homonym of prima Baranov (form of Wilhelmia equina), 
and a secondary homonym in Simulium s.l. of prima Baranov (19266, forms of aurea Fries, 
ornata Meigen and ruficornis Baranov). 

Wilhelmia stylata Baranov, form secunda Baranov, 19266 : 184, 186. Syntypes c?, Yugo- 
slavia: Macedonia, Treska (lost). 

The type-material of this form was preserved in alcohol and was lost in Vienna at the end 
of the war (Baranov i.litt. to Shewell). As Baranov mentioned the $ hypopygium (see dis- 
cussion under stylata above) there were presumably also slides of this structure, but these too 
are presumed lost. 

Rubzov (1959-1964) does not mention this name, but treats stylata as a synonym of 
Wilhelmia lineata (Meigen) ; by implication secunda is also a synonym of lineata. W. stylata 
i. secunda is a primary homonym of secunda Baranov (form of Wilhelmia equina), and a 
secondary homonym in Simulium s.l. of secunda Baranov (1926b, forms of aurea Fries, 
ornata Meigen and ruficornis Baranov). 

It should be noted that on p. 184 of the original publication the name secunda is mis-spelt 
'seunda' by typographical error, and that Smart (1945 : 513, 514) cites p. 185 for the original 
description in error. 



BIBLIOGRAPHY 

Note: The following bibliography contains a complete list of Baranov's papers 
on Simuliidae, although it has not been necessary to cite all of them in the foregoing 
text. Most of Baranov's papers are in Serbian (Cyrillic alphabet) or Croatian 
(Roman alphabet) and the treatment of the titles of these differs slightly in the 
following bibliography according to the language used : titles in Serbian are given in 
English translation only, titles in Croatian are given in the original and followed by 
an English translation in square brackets. Baranov used six languages in his 
papers and their summaries in various combinations, so we have thought it useful 
to append a note of the language (s) used in each publication. In most publications 
he used the V ending for his name, but occasionally used 'ff ' ; we cite all his papers 
under the spelling Baranov. For exactly half of the papers of which Baranov was 
author or co-author (17 out of 34 publications) there are short English abstracts in 
the Review of Applied Entomology, Series B (Medical and Veterinary) ; these provide 
helpful digests for many of the papers published in the rather difficult Serbo-Croat 
language, and we have cited the references to the R.A.E. abstracts (using this 
abbreviation) at the end of the relevant Baranov references. 



Babic, I., Baranov, N. & Ganslmayer, R. 1935. Die Kolumbatscher-Mticke im Jahre 
1934. Arch. wiss. prakt. Tierheilk. 69 : 205-212. [German: English abstract in R.A.E. 
(B) 23 : 275]. 



2i2 R. W. CROSSKEY & B. V. PETERSON 

Baranov, N. 1924. 'Golubachka mushitsa' (Simulium columbaczense sensu latiore.). Glasn. 

Minist. Poljopr. i Voda 2 (7) : 55-68. [Serbian with English summary: English abstract 

in R.A.E. (B) 13 : 50]. 

1925, with Radosavljevic. See Radosavljevic, D. M. & Baranov, N. 

1925. Neue Dipteren aus Serbien. Poljopr. ogled, kontr. Stanitsa Topcider. No. 1 : 1-11. 

[Serbian and German: English abstract in R.A.E. (B) 13 : 50]. 

1926a. Eine neue Simuliiden-Art und einige Bemerkungen iiber das System der Simulii- 

den. Neue Beitr. syst. Insektenk. 3 : 161-164. [German : English abstract in R.A .E. (B) 15 : 12]. 

19266. Uber die serbischen Simuliiden. Neue Beitr. syst. Insektenk. 3 : 183-194. [Ger- 
man: English abstract in R.A.E. (B) 15 : 78]. 

1927a. Guide for the identification of Simuliid larvae. Glasn. cent. hig. Zav., Beogr 

2 (3) : 93—96. [Serbian with German summary: English abstract in R.A.E. (B) 15 : 212]. 

• 19276. The larval development of some Simuliids. Glasn. cent. hig. Zav., Beogr. 2 (3) : 

97-104. [Serbian with German summary: English abstract in R.A.E. (B) 15 : 212]. 

1927c. Guide for the identification of Simuliid pupae. Glasn. cent. hig. Zav., Beogr. 

2 (4) : 91-93. [Serbian with German summary]. 

19270". Some morphological characters of the family Simuliidae and their importance 

in the classification of this family. Glasn. ent. Drust. [Acta Soc. ent. jugosl.] 2 : 19-25. 
[Serbian with German summary]. [Title only in R.A.E. (B) 16 : 192]. 

1928. On a biological method of control of the Golubatz fly. Jugosl. vet. Glasn. 8 (5) : 

137-138. [Serbian]. 

1929. O radnji Schonbauer-a, o golubackoj musici i o fauni srpskih simuliida [On the 

work of Schonbauer, on the Golubatz fly and on the Serbian simuliid fauna]. Jugosl. 
vet. Glasn. 9 (10) : 305-307. [Croatian]. 

1929, with Chorine. See Chorine, V. & Baranov, N. 

1934- Golubacka musica u godini 1934 [The Golubatz fly in the year 1934]. ^ et - Arh. 

4 (8-9) : 346-393. [Croatian with Russian summary: English abstract in R.A.E. (B) 

22 : 203]. 

J 935 fl - K poznavanju golubacke musice II. [Contribution to knowledge of the Golubatz 

fly II]. Vet. Arh. 5 (2-3) : 58-140. [Croatian with Russian summary: English abstract in 
R.A.E. (B) 23 : 161]. 

■ J 935^- Neues fiber die Kolumbatscher-Miicke. [Simulium columbaczense Schonb.) 

Arb. morph. taxon. Ent. Berl. 2 : 156-158. [German: English abstract in R.A.E. (B) 

23 : 275]. 

1935c. Die Kolumbatscher-Miicke im Jahre 1934. Arch. wiss. prakt. Tierheilk. 69 : 

205-212. [German]. 

1936a. K poznavanju golubacke musice III [Contribution to knowledge of the Golubatz 

fly III]. Vet. Arh. 6 (3-4) : 137-220. [Croatian with Russian summary]. [Title only in 
R.A.E. (B) 26 : 40]. 

19366. Studien an pathogenen und parasitischen Insekten IV. Simulium [Danubio- 

simulium) columbaczense Schonb. en Yougoslavie. Arb. parasit. Abt. Inst. Hyg. Zagreb 
No. 4 : 1-36. [French: English abstract in R.A.E. (B) 24 : 276]. 

1936c. Sadasnje stanje problema suzbijanja golubacke musice [The present position in 

the problem of Golubatz fly control]. Jugosl. vet. Glasn. 16 (10) : 524-527. [Croatian]. 

i937«- K poznavanju golubacke musice V [Studij epidemiologije golubacke musice 

na invaziji g 1936.] [Contribution to knowledge of the Golubatz fly V (Epidemiological 
study of the Golubatz fly in the year 1936]. Vet. Arh. 7 (5) : 229—276. [Croatian with 
German summary and descriptions: English abstract in R.A.E. (B) 25 : 249]. 

19376. Die Kolumbatscher Mficke in Jugoslawien im Jahre 1937. Arch. wiss. prakt. 

Tierheilk. 72 : 158-164. [German: English abstract in R.A.E. (B) 26 : 33]. 

1938a. K poznavanju golubacke musice VI (Studij golubacke musice i njezinih sinbio- 

cenonta) [Contribution to knowledge of the Golubatz fly VI (Study of the Golubatz fly 
and its synbiocenonts)]. Vet. Arh. 8 (7) : 313-328. [Croatian with German summary: 
English abstract in R.A.E. (B) 26 : 214]. 






BARANOV'S TYPES OF SIMULIIDAE 213 

Baranov, N. 19386. Die Kolumbatscher Miicke (Danubiosimulium columbaczense Schonb.). 

Z. hyg. Zool. 30 : 161-178. [German]. [Title only in R.A .E. (B) 26 : 224]. 
■ 1938c. Prilog poznavanju prirodnih neprijatelja golubacke musice iz klase insekta. 

[Contribution to knowledge of the natural enemies of the Golubatz fly from the class 

Insecta]. Arh. Minist. Poljopr. 5 : 106-115 (reprint pagination 1-12). [Croatian with 

German summary: English abstract in R.A.E. (B) 27 : 15]. 

1939a. K poznavanju golubadke musice VII (Bioloska svojstva golubacke musice i 

njezine seobe u g. 1938) [Contribution to knowledge of the Golubatz fly VII (Biological 
peculiarities of the Golubatz fly and its dispersal in the year 1938)]. Vet. Arh. 9 (3) : 105- 
125. [Croatian with German summary: English abstract in R.A.E. (B) 27 : 85]. 

19396. La mouche de Goloubatz. Bull. Off. int. Epizoot. 18 : 311-322. [French]. 

1939c Stand der Kolumbatscher Muckenforschung in Jugoslawien. Z. ParasitKde 

11 : 215-234. [German: English abstract in R.A.E. (B) 29 : 47]. 

19390". Odagmia ornata barensis subsp. nova i njen parazit Megaselia brevissima Schmitz. 

[Odagmia ornata barensis subsp. n. and its parasite Megaselia brevissima Schmitz]. Vet. 
Arh. 9 (11) : 599-601. [Croatian with German summary], 

1939c. Report of work on research and control of the Golubatz fly in the years 1934-1938. 

Publication of Moravian Ban's Administration, Nis [Yugoslavia], 48 pp. [Serbian]. [This 
is a printed and illustrated report issued by the regional governor (Ban) of Moravia, most 
of which was written by Baranov (as indicated in the foreword) although his name does not 
appear on the individual parts for which he was responsible. These, all in Serbian, are: 
pp. 21-28, 'Losses from the Golubatz fly'; pp. 29-34, 'Instructions for the collection of 
material' ; and pp. 35-48, 'Instructions for the observation of simuliids'. The work contains 
a coloured plate, between p. 42 and p. 43, of the male, female, pupa, eggs, and larva of the 
Golubatz fly.] 

1941- The Golubatz fly — on the history of the Golubatz fly. Poljopr. Glasn. [Novi 

Sad] 21 (5) : 2-4. [Serbian]. 

' 1942a. Boophthora argyreata Mg. u okolini Zagreba [Boophthora argyreata Mg. in the 

neighbourhood of Zagreb] . Vet. Arh. 12 (5) : 209-211. [Croatian with German summary]. 

19426. Die morderische 'Donau-Miicke'. Donauzeitung, issue 14th October 1942. 

[German: newspaper article on Golubatz fly including photograph of Danube river at 
Golubatz, figure of female of Golubatz fly, and map of Europe showing outbreak area, 
printed in German-language newspaper in German-occupied Yugoslavia.] 

& Nezic, Z. 1940. On the Golubatz fly in the year 1940. Account of laboratory work 

for investigation of the Golubatz fly at Golupcu for the year 1940. Jugosl. vet. Glasn. 
20 (11) : 410-414. [Serbian]. 

Chorine, V. & Baranov, N. 1929. Sur deux champignons parasites d' Anopheles maculipennis 
Mg. C. r. SSanc. Soc. Biol. 101 : 1025-1026. [French: English abstract in R.A.E. (B) 
17 : 247]. [Paper contains a reference to a fungus isolated from the larva of Simulium 
ornatum Mg.] 

Radosavljevic, D. M. & Baranov, N. 1925. Plant diseases and pests of the Timok valley 
[Serbia] in 1924. Work of the phytopathological section of the experiment and control 
station in Topcider. Glasn. Minist. Poljopr. i Voda, pts. 8-1 1 : 96-119. [Serbian]. 
[This paper is in two sections. The second section, beginning on p. 109, is entitled in 
Serbian 'II. Injurious insects of the Timok valley in the year 1924' and contains reference 
to the Golubatz fly on p. 109. The end of the paper contains a map (Map No. 2) of the 
distribution of Simuliidae in east Serbia.] 

Rubzov, I. A. 1956. Fauna SSSR (N.S.) No. 64, Diptera 6 (6) Simuliidae, 860 pp. [Russian]. 
Moscow & Leningrad. 

I 959-i964- Simuliidae (Melusinidae) in Lindner, Fliegen palaearkt. Reg. 14 : 1-689. 

[German] . 

1962. See 1959-1964 above. 

1963. See 1959-1964 above. 



2I 4 



R. W. CROSSKEY & B. V. PETERSON 



Sabrosky, C. W. & Crosskey, R. W. 1969. The type-material of Tachinidae (Diptera) 

described by N. Baranov. Bull. Br. Mus. nat. Hist. (Ent.) 24 : 27-63. 
1970. The type-material of Muscidae, Calliphoridae, and Sarcophagidae described by 

N. Baranov (Diptera). Proc. ent. Soc. Wash. 72 : 425-436. 
Sherban, E. 1961. New and little-known species of black-flies of the Eusimulinm aureum 

Fries group (Diptera, Simuliidae) from Rumania. Ent. Obozr. 40 : 677-685. [Russian 

with English summary]. [For English translation see Ent. Rev., Wash. 40 : 373-377]- 
Smart, J. 1945. The classification of the Simuliidae (Diptera). Trans. R. ent. Soc. Lond. 

95 : 463-532- 
Stone, A. 1963. An annotated list of genus-group names in the family Simuliidae (Diptera). 

Tech. Bull. U.S. Dep. Agric, No. 1284, 28 pp. 
Zivkovic, Vera. 1966. Les simulies (Diptera, Simuliidae) du groupe aureum en Yougoslavie. 

Acta vet., Beogr. 16 : 257-264. [Croatian with French title and abstract]. 



INDEX TO SPECIES-GROUP NAMES 

Note : Baranov's multiple uses of the infraspecific names prima and secunda are distinguished 
by giving the appropriate specific names in parentheses. 



agnatum, 200 
anderliceki, 196 
babici, 196 
barensis, 196 
bartulici, 196 
begbunaricum, 201 
borcici, 197 
brnizense, 201 
calopum, 203 
croatica, 195 
decolorata, 195 
djerdapense, 203 
echinatum, 194 
glumovoense, 204 
guelminoi, 197 
intermedia, 194 
kondici, 195 
litorale, 202 
mazedonica, 194 
nigrina, 195 
nikolici, 198 
ornatoide, 204 
prima (aurea), 104 



prima (equina), 206 
prima (ornata), 198 
prima (ruficornis), 199 
prima (stylata), 208 
profundale, 202 
pseudocolumbaczense, 204 
pukovacense, 204 
quarta, 207 
ruficornis, 199 
savici, 198 
secunda (aurea), 195 
secunda (equina), 207 
secunda (ornata), 198 
secunda (ruficornis), 200 
secunda (stylata), 208 
serbicum, 205 
stylata, 208 
tenuitarsus, 200 
tertia, 42 
tumanicum, 204 
vardaricum, 205 
zagrebiensis, 199 



Dr R. W. Crosskey, D.Sc, A.R.C.S., F.I. Biol. 
Commonwealth Institute of Entomology 
c/o British Museum (Natural History) 
London, SW7 5BD 




Dr B. V. Peterson, Ph.D. 
Entomology Research Institute 
Canada Department of Agriculture 
Central Experimental Farm 
Ottawa, Ontario 
Canada 



A LIST OF SUPPLEMENTS 

TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 
18 plates, 270 text-figures. August, 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 : 500 text-figures. September, 

I9 6 5- £3-25- 

5. Ahmad, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156: 
475 text-figures. November, 1965. (out of print) £2.15. 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. 

£3.I5- 

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 
text-figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera). Pp. 509. August, 1967. £8.50. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322 : 348 text-figures. August, 1967. £8. 

11. Mound, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp.172: 

82 text-figures. May, 1968. £4 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. 
November, 1968. £5. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198 : 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North-Western Europe 
(Hymenoptera : Chalcidoidea). Pp. 908: 686 text-figures. November, 1969. 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 
68 plates, 15 text-figures. October, 1971. £12. 



4S& 

•JXh Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 



REVISIONAL NOTES ON AFRICA' 






CHARAXES 
(LEPIDOPTERA : NYMPHALIDAE) 

PART VIII 






V. G. L. van SOMEREN 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 27 No. 4 

LONDON : 1972 



REVISIONAL NOTES ON AFRICAN 

CHARAXES 

(LEPIDOPTERA : NYMPHALIDAE) 

PART VIII 




BY 

VICTOR GURNER LOGAN van SOMEREN 

The Sanctuary, Ngong 
P.O. Box 24947, Karen, Kenya 



Pp. 215-264; 12 Plates, 7 Maps 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 27 No. 4 

LONDON: 1972 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 27, No. 4 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Ent.). 



Trustees of the British Museum (Natural History), 1972 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 29 December, 1972 Price £5 



REVISIONAL NOTES ON AFRICAN 

CHARAXES 

(LEPIDOPTERA : NYMPHALIDAE) 

PART VIII 

By V. G. L. van SOMEREN 



CONTENTS 



Synopsis .......... 

i. Charaxes hildebrandti Dewitz and its subspecies 

Systematic list ........ 

2. Charaxes thysii Capronnier, and C. hadrianns Ward and its 

subspecies ......... 

Systematic list ........ 

3. Charaxes imperialis Butler and its subspecies . 

Systematic list ........ 

4. Charaxes ameliae Doumet and its subspecies 

Systematic list ........ 

5. Charaxes lactetinctus Karsch and its subspecies 

Systematic list ........ 

6. Charaxes tiridates Cramer and its subspecies 

Systematic list ........ 

7. Charaxes fuscus Plantrou and C. numenes Hewitson and its 

subspecies ......... 

Systematic list ........ 

8. Charaxes bipunctatus Rothschild and its subspecies . 

Systematic list ........ 

9. The problem of Charaxes mixtus Rothschild, C. bubastis Schultz 

and related species ....... 

Systematic list ........ 

10. Further notes on Charaxes manica Trimen and its subspecies 

and Charaxes mccleeryi sp. n. . 

Systematic list ........ 

11. Further notes on the Charaxes xiphares complex and a descrip 

TION OF A NEW SUBSPECIES ...... 

Systematic list ........ 

Acknowledgements ........ 

References .......... 

Index ........... 



Page 
217 
218 
219 

219 
222 
223 
228 
228 
232 
232 
235 
235 
240 

240 

245 
246 
249 

249 

254 

255 
258 

259 
262 
262 
263 

264 



SYNOPSIS 

Fifteen species and their subspecies have been dealt with, two new species, seven new sub- 
species and three new forms have been described, and one name has been synonymized. 



2i8 V. G. L. van SOMEREN 

CHARAXES HILDEBRANDTI Dewitz and its Subspecies 

Charaxes hildebrandti (Dewitz) 

(PI. i, figs i, 2, Map i) 

Nymphalis hildebrandti Dewitz, 1879 : 200, T. 2, f. 16. 

Charaxes talagugae Holland, 1886 : 332, T. 8, f. 3. Type male; type-locality, Ogowe Riv., 

Gabon. 
Charaxes galba Distant, 1879 : 709. 

Male. Upperside. Resembling somewhat a small example of Ch. brutus angustus. Fore 
wing length 33-34 mm. Fore wing shape, apex slightly pointed, falcate, outer border slightly 
incurved at 3-4. Ground colour black, slightly brownish at base, where the veins are brownish. 
A creamy white bar crosses the wing from sub-apex to the hind margin, made up of five 
separated spots of increasing size from a small spot at costa to 5, the three upper spots arranged 
vertically, the spot in 4 set in a little, followed by spots of increasing size and conjoined; this 
bar is really made up of four postdiscal spots, the lower ones merging into the discal line and 
often represented by a glaucous blue border, frequently quite strong; thus the bar is often 
straight on the inner edge of the lower portion, the increase in size being on the distal side. 
Margin without any pale spots. Hind wing black, shading to greyish black on the inner fold ; 
the disc of the wing crossed by a creamy white bar, 3 mm at the costa then widening to space 2, 
where it tapers to and crosses the inner fold, above the anal angle, the expansion of the bar 
often glaucous blue, most evident on the outer border. No marginal spots, edge slightly 
dentate; tails short, 3-2 mm. Underside. Ground colour of both wings satiny greyish, with 
a more brownish tinge to proximal side of the white bars of both wings and to a lesser extent 
on the distal side, though strong in the curve of the fore wing. Base of fore wing costa whitish, 
shading to brownish beyond; cell with three sub-basal black dots and a wavy line beyond, 
with finer lines in sub-bases of ib-2, that in 2 often heavy. The inner edge of the white bar 
is accentuated by black lines from ia-3, the outer margin similarly bordered, separating the 
bar from a series of white and ochre lunules and represented in ib by a conspicuous round or 
angular black mark; beyond there may be a series of obscure darker submarginal marks, 
Hind wing ground colour as fore wing but basal area slightly darker, the dark zone of proximal 
side of white bar stronger; fine black lines in basal area limited to upper half; white bar distinct, 
greyish on distal edge, almost parallel-sided to ic, then narrowing and crossing the inner fold 
above the anal angle ; on its distal side there is a complete series of thin black lines accentuating 
the series of ochre-olive lunules which extend from the costa to the anal angle, where the 
olive expands and extends up the margin, internal to which is a series of lilac-whitish spots with 
black dots, double at anal angle; extreme edge black. 

Female. Fore wing length 39 mm. Upperside. General colour and pattern similar to 
that of male, but duller, the white bars of both wings wider. The ground colour of the fore 
wing is brownish black, slightly darker on the outer border. The creamy white bar similar in 
form to that of male; the upper postdiscal spots similarly separated to 4, the rest contiguous, 
but there is a slight indication of separation of the bar into two, from ib-4, by the presence of 
fine black lunules. Width of bar at ib-2, 4 mm, then narrowing. There is also an indication 
of whitish scaling in the discal line in 4-6. Hind wing ground colour brownish black in basal 
area, blacker on the border. The creamy white band is of about even width, 6 mm from costa 
to 2, then tapering and crossing the inner fold, the margins of the bar with greenish blue scaling, 
mostly apparent on the proximal side. The black border carries a series of obscure whitish 
marks from upper angle to hind angle. Margin of wing slightly dentate ; tails short and stumpy, 
4 and 3 mm. Underside. Ground colour of fore wing satiny bluish grey, somewhat like that 
of male ; the discal-postdiscal white bar clearly defined and with greater indication of separation 
of the two series of spots by crescentic black lines. The submarginal series of dark spots more 
visible, as also is the dark shading on the margin of the wing; the black tornal mark in ib 
distinct. Hind wing ground colour as in the male, the white band well defined; the series of 



. 



REVISIONAL NOTES ON AFRICAN CHARAXES 219 

ochre-olive lunules, black lined proximally, more distinct, so also the ochre-olive marginal 
border; extreme edge black. The submarginal series of whitish lilac marks accentuated by 
black distally, clearly visible. 

Range : Ashanti and Lagos to Cameroun, Gabon, N. Angola and western Congo. 



Charaxes hildebrandti katangensis Talbot 

(PI. 1, fig. 3, Map 1) 

Charaxes hildebrandti katangensis Talbot, 1928 : 229. 

Described from two males taken at Kinda, Katanga by F. G. Overlaet. Talbot 
states that these have the white bands of the upperside of both wings with only slight 
blue edging and the band of the hind wing slightly broader anteriorly. The under- 
side is pale with paler yellow postdiscal spots, which in the type are obsolete. 

Range: Apparently confined to Katanga, Congo (Kinshasa). 



SYSTEMATIC LIST 

Charaxes hildebrandti Dewitz 

Charaxes hildebrandti Dewitz, 1879. Type locality: Angola. 

Range: Ashanti, Lagos, Nigeria, Cameroun, Gabon, Bena Bendi, 

Angola, Congo (Kinshasa). 
Charaxes hildebrandti katangensis Talbot, 1928. Type locality: Katanga, Kinda. 

Range: Katanga, Congo (Kinshasa). 



2. CHARAXES THYSII Capronnier and C. HADRIANUS Ward and 

ITS SUBSPECIES 

Charaxes thysii Capronnier 

(PI. 1, fig. 4, Map 1) 

Charaxes thysii Capronnier, 1889 : 125, n. 70. 

Male. Fore wing length 32 mm. Shape not strongly falcate, but margin very bluntly 
dentate. Upperside. Fore wing, ground colour deep blue-black; a series of blue spots cross the 
wing in the postdiscal line, the three sub-apical in a row, elongate in shape, the upper one 
small, followed by a rounder spot in 4 set in a little, with a larger spot in 3 set at an angle to 
one above, so that the series is here kinked, the remaining spots of increasing size, more quadrate, 
that on the hind margin 4 mm. There are two somewhat obscured blue spots submarginally 
in ib with the faintest trace of spots in cellules above. Hind wing, ground colour blue-black 
shading to brownish grey on the inner fold; border of wing blacker. A blue bar crosses the 
wing from the costa to short of the hind angle, but represented on the inner fold by a whitish 
mark ; the costal spot does not reach the margin and is 4 mm in length, followed by larger marks 
in 5-6 projecting inward, the lower spots gradually taper to above the hind angle, thus the band 






V. G. L. van SOMEREN 




kzf 




REVISIONAL NOTES ON AFRICAN CHARAXES 221 

is almost straight on its outer margin and kinked on its inner, represented on the inner fold by a 
very small vertical streak. Admargin with a series of purplish blue lunules from region of the 
tails to 4 ; spots at anal angle greenish blue ; edge of wing bluntly serrate, with fine white fringe ; 
tails thin and sharply pointed, of about equal length, 5 mm, black with blue mid-line. Under- 
side. Fore wing ground colour a distinctive and very striking silvery white; the basal silvery 
area with three black spots in sub-base of cell, a zigzag line beyond and a black line at end of 
cell; heavier black lines sub-basally in ib and 2, followed by a slightly curved line through the 
sub-bases of 3-6, and continued in ib and 2 by broken lines. The discal line is crossed by a 
chocolate-brown bar, distally accentuated by a black line. Distal portion of wing with veins 
scaled with brown, the interspaces with less strong, long, arrow-shaped marks with points 
directed towards the marginal border, which is brownish, narrowly edged in black, the bases 
of the arrow-marks with silver-white marks adjacent to the postdiscal line; the tornal black 
marks are strong, double and elongate in ib and represented in 2 by a free spot. Hind wing, 
the basal silvery area crossed by an almost straight brownish line accentuated in black on 
proximal side and by black spots on distal edge, the black marks more lunate towards the anal 
angle, where the brown bar shades to olive; the basal area of the wing has a vertical, thin 'V 
mark, base toward costa and extending towards the lower arm of the cell, followed in turn by 
a thin, almost vertical line in the discal zone, running from the costa to above the anal angle, 
where it turns at right angles and crosses the inner fold. The border of the wing, distally to 
the heavy bar, has on its margin brick-red lunules accentuated proximally by black triangles 
shading to olive at the anal angle; edge of wing black with narrow white fringe. 

Female. This apparently is not known. 

Range : Originally described from the Kasai district of western Congo, the species 
has since been taken elsewhere in the Kelle area of the Moyen Congo, Congo 
(Brazzaville) . 

Charaxes hadrianus Ward 
(PI. 1, figs 5, 6) 

Charaxes hadrianus Ward, 1871 : 120. 
Charaxes dux Staudinger & Schatz, 1886 : 170. 
Charaxes gabonica Crowley, 1890 : 553. 

Male. Fore wing length 45-47 mm. Shape, apex bluntly pointed but outer margin not 
incised ; hind wing margin bluntly dentate. Upperside. Fore wing, base of wing and the bases 
of 1 a and 2 bright chestnut; the distal 2/3rd of the wing black, with a pattern of creamy white 
spots, those in the discal line of increasing size starting at the end of the cell with an elongate 
quadrate spot, followed by a larger quadrate spot at the sub-base of 3, a larger more angular 
mark with projection distad along 2, set in a little, then larger, more elongate in ib and ia, the 
ends shaded with bluish grey. Two small rounded spots present at about mid-point in 6 and 5 ; 
the spots in the submarginal line are small and rounded in the sub-apex, then larger in 3 and 2 ; 
the hind angle with an oval spot at end of ib. Hind wing creamy white with slight smoky 
suffusion at base, the disc of the wing immaculate, though the dark transverse bar of underside 
shows through. The submargin carries a series of black linear marks, heavier and tending to 
be conjoined in the region of the tails, a black mark at right angles present at the anal angle; 
margin with black portions extending up the tails; extreme edge white. Tails comparatively 
short, 5 and 3 mm long and sharply pointed. Underside. The basal portion of the fore wing, 
corresponding to the chestnut area above, is matt white shading to grey at the base of ib, where 
there is an elongate smoky brown streak; the distal portion of the wing is satiny silvery grey, 
varying with the direction of light, with the white spots of upperside faintly indicated, but the 
creamy areas of ia-ib stronger; a double black line indicates the end of the cell, and lesser 
variable black lines may be present in sub-bases of ib and 2. A conspicuous feature of the 



222 V. G. L. van SOMEREN 

underside is the large pear-shaped black spot in ib at the hind angle. Hind wing, ground 
colour white or silvery cream with a few discontinuous black lines crossing sub-bases of 3-7; 
the most striking feature is the chestnut bar, of almost even width extending from the costa to 
the inner fold at the anal angle, this bar though irregular on the distal edge, especially in 7, is 
accentuated by black edged with white. Submargin with a series of black dots corresponding 
in position to the black lines above, but here greyish; edge black, strongest on the tails. 

Range: Cameroun, Gabon, Congo (Brazzaville), Congo (Kinshasa), extending 
eastwards to the Semliki Valley. [For Map, see Part IX, to follow.] 



Charaxes hadrianus lecerfi Lathy 

(PI. 1, figs 7-9) 

Charaxes hadrianus lecerfi Lathy, 1925 : 94. 
Charaxes hadrianus Ward; Rothschild, 1900 : 400. 

Male. Fore wing length 45 mm. Upperside. Coloration and pattern very similar to the 
nominate race, but differing mainly in the increased size of the white spots in sub-bases of 5 and 6 
of the fore wing ; the increased size of the creamy white discal marks throughout, especially those 
from hind margin to 3, the increase in size being mainly on the distal border, so that the mark in 
2 is in contact with the larger white postdiscal spots in that area. There is, thus, a resemblance 
in the pattern to that of the female of the nominate race. Hind wing with slightly reduced dark 
basal area, but marginal dark border heavier, especially at the upper angle which encloses two 
white spots; these white spots visible and extending to the hind angle edged proximally in 
greyish; the submarginal black linear marks more contiguous, thus accentuating the marginal 
white border; edge black. Underside. Ground colour and pattern very similar to that of 
nominate race, distal portion of wing less shaded, but the black spot at the hind angle in ib 
strong. On the hind wing, the dark discal bar is slightly wider, less straight and tending to 
curve toward the inner fold above the anal angle ; the border is more greyish, so that the upper 
black spots are not so conspicuous. 

Range: Nigeria and Ghana; possibly extending westward to Sierra Leone where 
it may be represented by a separate subspecies. 



SYSTEMATIC LIST 

Charaxes thysii Capronnier 

Charaxes thysii Capronnier, 1889. Type locality: Kasai, S.W. Congo. 

Range: Western and Moyen Congo, Congo (Brazzaville). 



Charaxes hadrianus Ward 

Charaxes hadrianus hadrianus Ward, 1871. Type locality: Cameroun. 

Range: Cameroun, Gabon, Congo (Brazzaville), extending east- 
wards to Semliki Valley. 

Charaxes hadrianus lecerfi Lathy, 1925. Type locality: Warri, Nigeria. 

Range : Nigeria and Ghana and possibly extending to Sierra Leone. 






REVISIONAL NOTES ON AFRICAN CHARAXES 223 

3. CHARAXES IMPERIALIS Butler and its Subspecies 

The nominate race was described from the Gold Coast and Sierra Leone; other 
subspecies have been described from Cameroun, S. Congo and Malawi, thus the 
pattern of subspeciation follows that of many species with an east to west distribu- 
tion. 

Char axes imperialis imperialis Butler 

(PI. 2, figs 10-15, Map 2) 

Charaxes imperialis Butler, 1874 : 531, T. 11, f. 3; Butler, 1887 : 570, n. 26. 

Male. Fore wing length 40-48 mm; shape, apex rather pointed, outer margin slightly 
concave from 2-5, hind angle projecting slightly in ib. Upperside. Fore wing, ground 
colour blue-black; costa and basal veins brownish. Two small white subcostal dots in the cell 
with a larger one at its end; two small white spots in the upper discal line followed by a series 
of spots of increasing size from white spots in the sub-apex, those from 5 to hind margin blue, 
the mark in ib quadrate. Hind wing ground colour blue-black, slightly brownish at costa 
and shading to greyish black on the inner fold. Disc of wing bisected by a blue band of semi- 
detached spots, tapering slightly towards, but not encroaching on the inner fold, represented in 
the sub-costa by a white spot. Submargin with a series of blue triangular spots, double at 
anal angle; admargin with interrupted blue lunules, the blue extending up the tails which are 
short and sharply pointed, 4 and 2 mm long. Underside. Brownish grey with a slight olive- 
ochreous tinge. Pattern not strong, consisting of three thin black transverse lines in the cell, 
the middle one divided into two spots, all outlined in white, a straighter black line at end of 
cell, and sub-basal black lines in 4 and 2 and an angled mark in ib. In the discal line there are 
small whitish triangular marks followed by thin black lines in ib-3. In the postdiscal lines 
are whitish spots, corresponding to the blue spots of upperside, distally shaded with olive- 
ochreous, the mark in ib lined with black proximally and with a bold black horse-shoe spot 
distally and with a small black spot in space above, forming a conspicuous 'eye-spot'. Margin 
of wing slightly shaded with olive-ochreous at hind angle. Hind wing ground colour as fore; 
basal area crossed by thin black lines outlined in white; a similar series in the discal line, followed 
by a row of whitish lunules outlined in black proximally and shaded with olive-ochreous distally, 
the mark in the anal angle stronger; the submargin with a row of whitish lilac linear marks with 
black dots distally; border olive-ochreous with white shading at end of veins, edge black with 
narrow white fringe; anal angle with twin black spots. 

Female. Fore wing length 45-48 mm. Upperside. Ground colour of fore wing brownish 
black with blue sheen at base. A crescentic white mark at end of cell and a series of larger 
whitish spots in the discal line from sub-costa to 2, represented on the hind margin in ia-ib 
by a blue quadrate patch, linking up in these areas with the postdiscal series of bufnsh ochreous 
spots of decreasing size, curving toward the costa in sub-costa in 5-7. Margin with a series of 
bufnsh ochreous spots, double at hind angle, decreasing in size to apex. Costa of wing brown. 
Small whitish spots are sometimes present in the subcostal region of the cell. Hind wing 
ground colour darker than fore wing, especially on the border. Disc of wing with a blue patch, 
sometimes extending basad in the cell and represented in the subcosta by one large white spot 
and a small buffy spot more distad. Submargin of wing with huffish ochreous spots decreasing 
in size from upper angle to anal angle where the spots are double. Margin of wing with inter- 
rupted huffish ochreous lunules; edge black. Tails thin and pointed, upper 6 mm, lower 4 mm. 
Underside. Ground colour as in the male, sometimes of a colder greyer tone; pattern similar 
to that of male, the fore wing spots rather more prominent. Hind wing pattern as in the male. 

Range: Sierra Leone, Ivory Coast, Ghana, to western Nigeria; ? Liberia. 



224 



V. G. L. van SOMEREN 










• OH n ♦-$£<<$ 



REVISIONAL NOTES ON AFRICAN CHARAXES 225 

Charaxes imperialis albipunctus Joicey & Talbot 

(PI. 3, figs 16-21, Map 2) 

Charaxes imperialis albipuncta Joicey & Talbot, 1921 : 68. 

Joicey and Talbot based this race mainly on the fact that female specimens from 

Cameroun had white spots on the fore wing. As I have no topotypical examples 

from the Ja River at Bitje, I take the liberty of quoting the original description: 

'Male, upperside of fore wing not constantly different from typical form. 

Hind wing with a broader band in most specimens. The inner spot in cellule 7 

is blue or only slightly white. Underside of fore wing with the yellow proximal 

border to the black submedian spot edged with blue, and this again with black 

proximally. Hind wing with the postdiscal bar in cellule 7 either without white 

distal scaling or with only a trace of it. Female, upperside of fore wing with 

the spots white and smaller than in typical form. The black inner marginal 

band is reduced anteriorly and does not reach vein 2. Hind wing with the 

band not reaching vein 7. The outer spot in cellule 7 smaller and without any 

brown tinge. Submarginal and marginal spots white. Underside as in the male. 

Fore wing with spots as above. No spot bordering the submedian bar. Hind 

wing with postdiscal spots further from the discal bars than in typical form.' 

No measurements are given. 

One is well aware of the fact that within the area termed theCameroons (Cameroun), 
there are many species with a restricted distribution, but nevertheless, it is of 
interest to note that in the case of Ch. imperialis, female specimens from the border 
between Cameroun and Congo (Brazzaville), at Ouesso, are not white-spotted, nor 
are the females from eastern Nigeria, at Ikom. It is of interest to note that Talbot 
places males from the Beni-Ituri forests of eastern Congo with albipunctus, but 
whether this is supported by white-spotted females he does not say. It is also 
worth noting that males from the adjacent Semliki River at Bwamba Valley are 
similar to males from eastern Nigeria and western Congo examples, but the females 
are not white-spotted, and they differ markedly from examples of ssp. ugandicus 
ssp. n. from the western shore of Lake Victoria at Katera. 

The probability is, that if females in Cameroun are always white-spotted, the 
race albipunctus is limited to that area only and does not extend east to Uganda! 



Charaxes imperialis ugandicus ssp. n. 

(PI. 3, figs 23, 24, Map 2) 

Charaxes imperialis albipuncta Joicey & Talbot; van Someren, 1935 : 12, T. 25. 

We are faced with an additional problem in Uganda, for as already stated, the 
males from the Bwamba Valley east of the Semliki River, are very similar to males 
from eastern Nigeria. The white spots in the basal half of the fore wing may be 
absent or limited only to a minute dot at the end of the cell, or the discal spots may 



226 V. G. L. van SOMEREN 

be present. The postdiscal row of blue spots is very similar to that of east Nigerian 
examples, the subcostal sub-apical spot may be white or blue. The hind wing blue 
patch is broader than central Uganda examples. 

Thus Ch. imperialis in Uganda presents an interesting problem, not only from the 
point of view regarding males, but also the females. Bwamba females are smaller 
than those from Katera on the west shores of Lake Victoria, with a fore wing length 
of 47-50 mm. The fore wing ground colour is browner, the spots are smaller and 
the postdiscal spots are not as rich orange as in Katera examples; the blue patch 
on the hind margin is more broken up. The hind wing blue patch is restricted; the 
white spot on the costa is large, the outer one usually a dot. The submarginal row 
of yellow-ochre spots may be small or large. The underside pattern may be reduced 
and faint or strong. Tails thin and pointed, 5 and 3 mm. 

Male. Fore wing length 45-52 mm. Shape similar to other races. Upperside. Fore 
wing, ground colour blue-black with strong greenish blue sheen at base and over the cell. Pat- 
tern similar to that of other races ; spots at end of cell and in discal line white and larger than in 
other subspecies; the series of spots in the postdiscal line bright blue, the marks in ia-ib (par- 
ticularly the latter) narrow, that in 2 ovoid, the spots above and in sub-apex small and mostly 
whitish. Hind wing, ground colour as fore wing, the outer border wider than in other races, 
due to a reduction in the width of the blue, which does not extend so far toward the inner fold ; 
the inner spot in 7 is large and white, that beyond, blue. The submarginal series of blue spots 
well developed; the interrupted admarginal blue lunules usually extending to 6. Tails very 
short, upper, 2 mm. Underside. Ground colour clay-brownish grey, with a slight satiny 
sheen mainly in the curve of the fore wing and border of hind wing. An occasional specimen 
has a more brownish tone. The rather sparse markings are subdued, but similar in distribution 
to that of other races; the conspicuous feature is the ocellus at the hind angle of the fore wing. 

Female. This is noted for its large size and strong coloration; length of fore wing 52-57 mm. 
Shape like the male but hind wing more rounded at hind angle. Upperside. Fore wing, 
ground colour brownish black with a purplish sheen at base. The white spot in the cell end is 
round, the two in the upper discal line angled, those in sub-bases of 2 and 3 angled or ovoid; 
spots in the postdiscal row are bold tawny orange in colour, the largest being in 2 and 3, that in 
ib small and fused with the blue patch in ia-ib, the whole patch rather narrow, the longest 
mark being in upper part of ib. Margin of wing with distinct orange triangles with a fringe of 
white on edge opposite each spot. Tails rather short, upper 6 mm, lower 3 mm. Underside. 
Ground colour clay-brownish grey or paler ochreous grey. The black wavy transverse lines 
in the cell are thin, and edged with white; those in sub-bases of ib and 2 heavier, the whitish 
spots in the discal line are outlined in black proximally; the postdiscal series are represented by 
lilac white lunules strongly shaded ochreous distally and edged in black; the tornal black loop 
mark in ib is strong, distally shaded with whitish. Hind wing, ground colour as fore wing; 
the basal black lines thin, those in the discal line offsetting a series of whitish lunules or just 
whitish shading distally; the postdiscal sinuate line of narrow lilac-white lunules, shaded 
ochreous olive distally, are edged internally in black, strongest at the anal angle. The sub- 
margin carries a complete row of lilac -white spots, not sharply defined, but those in the region 
of the tails with a black spot distally, double at anal angle ; border of wing with ochreous lunules 
shaded whitish distally ; anal angle with olive ground ; edge of wing somewhat darker greyish, 
narrowly fringed with white; tails outlined in greyish olive as edge. 

Holotype <$. Uganda: Katera Forest, Masaka District, west shore of Lake 
Victoria (van Someren), in B.M.(N.H.). 

Allotype $. Uganda: Katera Forest, Masaka District (van Someren), in 
B.M.(N.H.). 



REVISIONAL NOTES ON AFRICAN CHARAXES 227 

Range: Uganda, central and western; Katera Forest, west shore of Lake Victoria, 
Budongo Forest, Bugoma Forest. Kibale Forest, Toro. Local variation, Bwamba 
Valley, east side of Semliki River. 

$ f. caerulipunctus forma n. 
(PI. 3, fig. 22) 

Some Bwamba females have the discal and postdiscal spots of the fore wing blue 
in colour and bold (Bwamba Valley, Grahame Coll.). A blue spotted variation is 
also found amongst these examples. The hind wing blue patch is rather broken up, 
but the submarginal and marginal marks are bold. 

Charaxes imperialis paulianus Rousseau-Decelle 

(PI. 4, figs 25, 26, Map 2) 

Charaxes imperialis pauliana Rousseau-Decelle, 1933 : 269. 

Male. Fore wing length 45 mm. Shape as in other races. Upperside. Fore wing ground 
colour blue-black; costa brownish. Pattern of blue spots similarly arranged to that of other 
races, the row of postdiscal spots slightly stronger blue, the spot in the cell white, those beyond 
in the discal line tinged with blue as are the small upper spots in the postdiscal series. Hind 
wing, ground colour and pattern as in other races, the hind end of the blue patch fading out 
somewhat as it nears the inner fold. The submarginal spots strongly blue; the broken marginal 
border blue but limited almost entirely to the region of the tails and hind angle. Underside. 
Ground colour rather browner than in other races, but pattern very similar; there is, however, a 
dark triangular subcostal spot beyond the end of the cell. The hind wing ground colour 
slightly darker, the pattern rather obscured except for the subcostal whitish spot in discal 
line, and the submarginal pale spots; the blackish sub-basal spot in 5 more distinct. 

Female. Fore wing length 50 mm. Upperside. Fore wing, ground colour and pattern 
as in other races but the blue mark rather reduced, so also that of the hind wing but the white 
costal spot in the discal line large; the submarginal spots also large. Underside. This reflects 
the reduction in the blue areas but in the fore wing the spotting is well developed; this also 
applies to the hind wing, the submarginal row of light spots being a strong feature. 

Range: So far recorded only from the Katanga area of S.E. Congo (Kinshasa). 

Charaxes imperialis ludovici Rousseau-Decelle 

(PI. 4, figs 27, 28, Map 2) 

Charaxes imperialis ludovici Rousseau-Decelle, 1933 : 211. 

Male. Fore wing length 45 mm. Upperside. Fore wing, ground colour blue-black with 
greenish sheen at base, costa chestnut-brown. Spot at end of cell and upper discal row white and 
distinct, the postdiscal blue spots whitish and slightly angular, the mark in ib somewhat narrow, 
that in ia extended distad. Hind wing blue patch rectangular and extending slightly into the 
inner margin, the two costal marks large and distinct; the submarginal blue spots large; the 
upper tail longer than in other races, 6 mm, that at hind angle, 2 mm. Underside. Ground 
colour paler than in other races, but markings similar; on the hind wing the postdiscal sinuous 
line of pale lunules with darker shading distally so that the line shows up more clearly. 

Female. Fore wing length 48 mm. Upperside. Fore wing, ground colour brownish 
black, paler and more olive toward the base and along the costa ; pattern of light spots arranged 



228 V. G. L. van SOMEREN 

as in females of other races but the blue patch at the hind margin connecting up the discal and 
postdiscal spots in that area reduced in size, especially that in ia; the discal spots and that at 
end of cell whitish, those in the postdiscal row ochreous. The outer margin with well developed 
creamy spots extending from the hind angle to the apex. Hind wing, ground colour blacker 
than fore especially on the broad border of the wing. The blue patch not sharply defined, its 
distal border 'toothed' and dull blue, not reaching the inner fold which is ashy grey. The 
submarginal row of ochreous spots strong, the two spots in the subcosta large and whitish; 
the interrupted marginal border strongly marked. Tails longer and sharply pointed, upper 
8 mm. Underside. Ground colour slightly darker than that of male; pattern of light spots in 
the fore wing slightly stronger, that of the hind wing equally indistinct, the ocellus in the hind 
angle of the fore wing being a strong feature. 

Descriptions taken from a pair captured on the Lisombe River, Zambia (C. B. 
Cottrell). These are, perhaps, not quite typical since the type specimens are from the 
north-eastern area of Malawi (Nyasaland) (teste Gifford). 

Range: Malawi and adjacent Zambia. 

SYSTEMATIC LIST 

Charaxes imperialis Butler 

Charaxes imperialis imperialis Butler, 1874. Type localities: Sierra Leone( <$), 

Ghana ($). 

Range : Sierra Leone, Ghana, Old Calabar, ? Nigeria. 
Charaxes imperialis albipundus Joicey & Talbot, 1921. Type locality: Bitje River, 

Cameroun. 

Range: Cameroun and ? Northern Congo. 
Charaxes imperialis paulianus Rousseau-Decelle, 1933. Type locality: S. Congo, 

Katanga, Kafakumba. 

Range: S. Congo, Katanga Pro v. 
Charaxes imperialis ludovici Rousseau-Decelle, 1933. Type locality: Lake Nyassa. 

Range: Malawi, L. Nyassa and adjacent Zambia. 
Charaxes imperialis ugandicus ssp. n. 

Range: Uganda, west shore of L. Victoria, Katera Forest; 

Budongo and Kibali forests in western Uganda. 
Charaxes imperialis $ f . caerulipunctus forma n. Type locality : Bwamba, Uganda. 

Range: Variations in the Bwamba Valley, east side of Semliki 

River. 

4. CHARAXES AMELIAE Doumet and its Subspecies 

Charaxes ameliae ameliae Doumet 

(PL 4, figs 29, 30, Map 3) 

Charaxes ameliae Doumet, 1861 : 171, T. 5, f. 1. 

Charaxes ameliae Doumet; Hewitson, 1876 : T. 49, f. 21-22. 

Charaxes regius Aurivillius, 1889 : 191. 

Male. Fore wing length 46-48 mm. Upperside. Fore wing, ground colour deep blue- 
black; pattern limited to a series of blue spots in the subcostal region and postdiscal line as 



■a 



REVISIONAL NOTES ON AFRICAN CHARAXES 



229 




230 V. G. L. van SOMEREN 

follows: an elongate mark at the upper part of the base of the cell, often with a black dot just 
short of the centre; some blue scaling along the costal vein; a quadrate spot at the end of the 
cell, and three blue marks, the middle one largest, in the sub-base of 5-7. A postdiscal series 
of blue spots increasing in size from subcosta in sub-apex to the hind margin, the upper four in a 
curve, that in 3 set in a little, that on the hind margin sometimes a streak, there is also a blue 
streak in the lower basal area of ib. Margin with a series of small blue spots, double in ib. 
Hind wing ground colour blue-black shading to ashy grey on the inner fold ; disc of wing with a 
crescentic blue mark from base of 3 and the end of cell and tapering at base of 3, represented 
on the subcosta by a separate blue spot. Margin of wing with a series of rounded blue spots, 
double at anal angle; marginal border with blue lunules. Edge bluntly dentate, tails short and 
robust, upper 3-4 mm, lower 2 mm. Underside. Ground colour brownish grey with olive 
tinge, paler at the base of the cell and bases of 1 and 2, somewhat variable. Basal black marks 
strong, three black marks at basal half of cell followed by a thin wavy line and a thinner one at 
end of cell, and stronger black lines sub-basal in ib and 2, followed by a series of curved thinner 
lines on inner discal line, those in ib joined by a black line, these spots and lines outlined in 
white. In the postdiscal line is a series of spots of increasing size, the three upper ones small, 
with some dark shading distally, the lower spots large, that at the tornus and space above, 
black-centred and outlined in black proximally forming conspicuous 'eye-spots'. Marginal 
edge narrowly white. Hind wing ground colour as fore wing, with a paler bar in sub-basal 
area, thinly outlined in black and extending from the costa to short of the inner fold, followed 
by a discal pale bar outlined proximally in black. In the postdiscal line a series of whitish 
lunules, that in 5 set well in and distally shaded with darker colour ending at the anal angle 
in a double thin black crescentic mark. The submargin with a complete series of pale whitish 
spots with slight black distally ending in the double back dots in the anal angle. Marginal border 
brownish grey, edge very narrowly white. There is some slight variation in the spotting on the 
upperside, mainly in regard to the width of the blue patch in the disc of the hind wing. 

Female. Fore wing length 46-52 mm. Shape as in the male. Upperside. Fore wing, 
ground colour umber-brown, shading into black at end of cell, outer half of wing black. There 
is a large semi-quadrate white spot just beyond the end of the cell, and beyond, in the discal 
line two elongate spots and a streak in subcosta above ; no spot in 4 but those in 3 to the hind 
margin increasing in size, the mark in ia is 8 mm. All these spots are creamy white. Post- 
discal row of creamy white spots complete from sub-costa to ib. Marginal internervular spots 
clear and distinct, of increasing size from apex to the double spot in ib. Extreme edge very 
narrowly white, opposite the spots. Hind wing, ground colour at base umber-brown shading 
to ashy grey on inner fold. Border of wing black; disc crossed by a white band, 8 mm wide at 
the costa, rapidly decreasing in width, ending in a line which crosses the inner fold above the 
anal angle, the inner edge of this band is straight but with some bluish white scaling over the 
end of the cell, while the outer edge is irregular with toothed projections in the mid area to the 
costa, ending in two discrete white spots in 5-7. Submarginal series of round spots, white and 
large at the upper angle, decreasing in size and bluish in colour opposite the tails, ending in the 
double smaller spots of the anal angle. Admarginal row of white lunules complete, becoming 
bluish at the anal angle. Margin bluntly dentate, tails thin and sharply pointed, upper 6 mm, 
lower 3 mm. Underside. Ground colour as in the male, but pattern more contrasty, black 
marks at base of fore wing stronger, those in the proximal edge of the white discal bar strong. 
Postdiscal series of white spots as above but more strongly outlined in black proximally, and 
distally with a series of whitish ocelli with dark centres, very strong at the tornus and space 
above, less distinct above, but extending up to the sub-apex. Margin with pale spots of decreas- 
ing size from ib to apex. Hind wing ground colour more brownish olive-grey to inner edge of 
white discal bar, which is crossed by a white vertical line, outlined blackish in 7-6, through the 
cell and sub-base of ic. Discal white bar as above, clear-cut and almost straight on the inner 
edge, more dyslegnic on outer border. In the postdiscal line are whitish spots and crescentic 
marks distally shaded with diffuse olive-brown. Triangular white spots present on the sub- 
margin with black spots distally in region of tails and separated from the marginal whitish 
lunules by an ochreous zone, turning olive at the anal angle. Edge black. 



REVISIONAL NOTES ON AFRICAN CHARAXES 231 

The above descriptions are taken from specimens collected in the Ivory Coast. 

Range: Sierra Leone, Liberia, Ghana, Ivory Coast, Nigeria, Cameroun to northern 
Congo. 

Charaxes ameliae victoriae ssp. n. 

(Pis 4, 5, figs 31-36, Map 3) 

Rothschild gives the measurements of West African specimens as <$ 38-44 mm, 
$, 42-54 mm. A long series from Uganda are noticeably larger, the fore wing 
measurements are <$ 47-50 mm, the majority 50; $ 51-58 mm, majority 57 mm. 

This is in keeping with the large majority of species with a west to east distribution, 
the eastern aggregates being generally the larger. As in many, the difference in 
size is accompanied by constant differences in coloration and pattern in one or 
both of the sexes. 

Male. Upperside. Ground colour as in the nominate race, deep blue-black, the pattern of 
blue spots similarly arranged but larger and bolder. Underside. Pattern as in nominate 
ameliae, but ground colour not so dark, less brownish thus the pattern is not in such contrast. 

Female. Upperside. Larger than the nominate race, the fore wing spots larger, creamy to 
ochreous in colour; the marginal spots in the fore wing larger. The marginal and submarginal 
spots in the hind wing more prominent. Underside. Ground colour less dark, but pattern 
bolder owing to increased size of light markings. 

Holotype male. Uganda : Katera Forest, west shore Lake Victoria {van Someren), 
in B.M.(N.H.). 

Allotype female. Uganda: Katera Forest, west shore, Lake Victoria {van 
Someren), in B.M.(N.H.). 

Range: Uganda, from the Semliki Valley and western Uganda, including the 
south-west, to Busoga in the east. 



Charaxes ameliae amelina Joicey & Talbot 

(PI. 5, figs 37-39, Map 3) 

Charaxes ameliae amelina Joicey & Talbot, 1925 : 644. 

This 'race' was based on the fact that in Malawi all the females are white-spotted, 
not creamy. Attention was drawn to this fact by Rothschild, 1900, who however 
pointed out that 'white-spotted' females also occurred in the western aggregate, 
but did not state in what proportion ; moreover his 'nominate race' embraced speci- 
mens from Sierra Leone to the Niger and Congo in the region of Beni. I have already 
drawn attention to the fact that specimens from western Africa are smaller than 
specimens from Uganda; moreover, in a series of 16 examples from Uganda, only 
one has the spotting of the fore wing white, the rest are creamy or even ochreous. 
Malawi specimens are larger than west African examples. On this evidence I 
support the recognition of the 'race' amelina. 



232 V. G. L. van SOMEREN 

Male. Fore wing length 48-50 mm. Upperside. Fore wing, ground colour deep blue- 
black, with a strong blue sheen; pattern of blue marks on both wings bolder than in nominate 
race and more like those of the Uganda subspecies victoriae. Underside. Even paler than 
that of the Uganda subspecies, and much paler than nominate. 

Female. Fore wing length 48-52 mm. Upperside. Fore wing, ground colour as in other 
subspecies, but spotting of the fore wing as large as in the subspecies victoriae, that of the hind 
wing resembles more that of the nominate, the spotting on the border being smaller. Underside. 
Very similar to ameliae victoriae. 

Range: Malawi, mostly in the forests of the Nkata Bay area and the Nyika; also 
recorded from the Mwinilunga area of Zambia. Specimens from the Kigoma area, 
N.E. of Lake Tanganyika, belong to this subspecies. There is no record from 
Katanga, but Schouteden records 'ameliae' from the Belgian Congo without an 
exact locality. 

SYSTEMATIC LIST 

Charaxes ameliae Doumet 

Char axes ameliae ameliae Doumet, 1861. Type locality: Gabon. 
Synonym. Charaxes regius Aurivillius, 1889. Kamerun. 

Range : Sierra Leone to upper Congo. 
Charaxes ameliae amelina Joicey & Talbot, 1925. Type locality: Malawi. 

Range: Malawi, N. Zambia, ? Katanga. 
Charaxes ameliae victoriae ssp. n. 

Range: The Semliki Valley and western Uganda, including the 

S.W. area to Busoga in the east. 



5. CHARAXES LACTETINCTUS Karsch and its Subspecies 

Charaxes lactetinctus Karsch 

(Pis 5, 6, figs 40-42, Map 4) 

Charaxes lactetinctus Karsch, 1892 : 113. 

Male. Fore wing length 38 mm; shape, apex rather pointed, outer margin concave in 3-4, 
hind angle projecting. Upperside. Fore wing, base from lower part of cell to mid hind margin 
bluish white, distal portion of wing black with tawny red spots in pattern from base of costa 
and upper part of cell, a tawny red spot at base of 4, and two quadrate spots beyond linking up 
with the series of tawny red spots in the postdiscal line, which extends from the costa to 2, 
sometimes represented in ib by a small spot; border of wing from hind angle to apex tawny 
red. Hind wing, basal area bluish white, shading to greyish ochre on the inner fold; a large 
diffuse orange spot at mid costa separating the upper part of the bluish white base from the 
broadly black border ; marginal border tawny rufous from upper angle to upper tail ; submargin 
with greyish blue spots, large and distinct in region of tails, but small above, the mark on the 
anal angle more lilac, with white dots; margin at anal angle olive. Edge of wing black with 
slight white fringe to upper tail. Tails thin and pointed, 4-5 mm long, the lower tail fused at 
the base with the projecting anal angle. Underside. Ground colour purplish chestnut, especial- 
ly on hind wing. The fore wing chestnut shades to purplish grey at bases of ib and ia. The 
underside pattern is remarkably scanty, consisting of two silver and black lines in the sub-base 
of the cell, ib and 2, with bolder silvery line toward the end of the cell, slightly outlined in black. 
In the discal line at the costa is a triangular silvery mark, widest at 8 and tapering to 5 ; there 



REVISIONAL NOTES ON AFRICAN CHARAXES 



233 




234 v - G - L - VAN SOMEREN 

is also an indication of the rufous spots of upperside, enhanced on either side in ib- 3 by black 
marks on the proximal side, and edged with black and bluish grey on the distal side in ib, 
lessening in 2—3 and faintly indicated in spaces above. Hind wing almost immaculate in the 
disc, there are two small white spots in 8; in the postdiscal line there is an obscure continuous 
darker bar from costa to above the anal angle beyond which the border is more tinged with 
lilac, culminating in the distinct lilac edge to the rufous border, which has a series of greenish 
lines with central black dots, the black spots accentuated in the anal angle by the olive ground. 

Female. Larger than the male, fore wing length 44 mm. Upperside. General colour and 
pattern resembling that of the male, but bolder. The bluish white areas of fore and hind wing 
slightly more extended. 

Range: The nominate race extends from Ivory Coast to Nigeria and Cameroun, 
Central African Republic, Congo (Kinshasa), Congo (Brazzaville), but is not yet 
recorded from Katanga. 



Charaxes lactetinctus busogus ssp. n. 

(PI. 6, figs 43-46, Map 3) 

Shape similar to western examples but slightly larger than the nominate race. 

Male. Fore wing length 42-43 mm. Upperside. Coloration and pattern very similar, 
but bluish white basal areas slightly more extended on both wings, the black border of the hind 
wing thus reduced in width. Underside. Pattern bolder, especially that of the hind wing. 

Female. Upperside. Basal bluish white areas more extended on both wings, the pattern 
bolder. Underside. Ground colour paler, but pattern more distinct and bolder. 

Holotype male. Kenya: Busoga district, Broderick Falls Scarp (van Someren), 
in B.M.(N.H.). 

Allotype female. Uganda: Metu Hills, N.W. Madi, West Nile, v-vi.1954 (van 
Someren), in B.M.(N.H.). 



$ form jacks onianus van Someren 

(PL 6, figs 47, 48) 

Charaxes lactetinctus i. jacksonianus van Someren, 1936 : 174. 

Upperside. Differs from the normal female in having the basal bluish white areas suffused 
over with tawny orange, especially in the region of the fore wing cell. The pattern of the fore 
wing generally larger and bolder, but paler ; the black areas not so dark. On the hind wing the 
bluish white area is slightly suffused over with tawny orange especially at the mid-costa and 
along the distal border; the black border is not so dark. Underside. Ground colour of fore 
wing not so dark as usual, thus the pattern stands out more boldly; the black and silvery lines 
in the cell more developed. The pattern of the hind wing bold on a paler ground, and there is 
more silvery white in the basal area of the costa. 

The type specimen was taken in the TransNzoia district of Kenya. 

Range: Uganda, mainly in the central and eastern districts, and also found on 
the west side of Lake Rudolf and West Nile Districts, extending to the northwestern 
area of Kenya. 



REVISIONAL NOTES ON AFRICAN CHARAXES 235 

Charaxes lactetinctus ungemachi Le Cerf 

(PI. 7, fig. 50, Map 3) 
Charaxes lactetinctus ungemachi Le Cerf, 1927 : 144. 

Described by Le Cerf from a male and two females from Youbdo (Birder), Ethio- 
pia. The paratype female figured here well illustrates the subspecific differences 
noted by Le Cerf, which makes the repetition of the original description superfluous. 



<$ form brunneus Carpenter 
(PL 7, ng. 49) 

Charaxes lactetinctus ungemachi cJ f. brunneus Carpenter, 1935 : 359. 

Fore wing length 36 mm. Upperside. Fore and hind wing basal areas mahogany-brown, 
without any trace of the bluish white usual in these areas in normal specimens; the brown of 
the hind wing merges into the black border. 

Range: Ethiopia, Youbdo. 



SYSTEMATIC LIST 

Charaxes lactetinctus Karsch 

Charaxes lactetinctus lactetinctus Karsch, 1892. Type male in B.M.(N.H.). Type 

locality: Togo. 

Range: Ivory Coast, Ghana, Togo, Cameroun, Central African 

Republic, Congo. 
Charaxes lactetinctus busogus ssp. n. 

Range: Uganda, Tororo, Majanji; N.W. Kenya: Broderick Scarp, 

Kabras, Busia, Saboti Hill, Kitale, TransNzoia, Lugari, Kitesh. 
Charaxes lactetinctus jacksonianus $ form, van Someren, 1936, TransNzoia, S.E. 

Mt. Elgon. 
Charaxes lactetinctus ungemachi Le Cerf, 1927. 
Charaxes lactetinctus ungemachi <$ f. brunneus Carpenter, 1935. 

Range : Ethiopia, S. W. ; Haete River. 



6. CHARAXES TIRIDATES Cramer and its Subspecies 

Examination of a considerable series of tiridates, from various areas of its distribu- 
tion from West Africa to East Africa, indicates that there is a considerable difference 
in the appearance in the populations in the two extremes, with transitionals in the 
intervening countries. I shall deal with these differences in the following arrange- 
ment. 



236 V. G. L. van SOMEREN 

Char axes tiridates tiridates (Cramer) 
(PI. 7, figs 52, 54, Map 5) 

Papilio Eques Achivus tiridates Cramer, 1777 : 100. 

Char axes tiridates (Cramer); Doubleday, 1844. [Corrected type-locality: Ashanti, Gold Coast]. 

Char axes marica Fabricius, 1793 : 113. [Locality: 'Africa'. ? specimen]. 

Charaxes marica Fabricius; Watkins, 1923 : 209. 

Male. Fore wing length 45-47 mm. Upperside. Fore wing, ground colour black, browner 
on the costa. A series of small blue spots in the upper median or discal line, not very strong, 
extend from the subcosta, just beyond the end of the cell, as a streak and a rounded spot, 
followed by two spots, set slightly out in sub-bases 3 and 2. Postdiscal spots, complete, upper 
one in subcosta small and white followed by blue spots in 6-5 set out slightly, spots in 4-3 
set in, followed by a blue spot in 2 and double spot in ib, occasionally a spot in ia towards the 
hind angle. Marginal golden lunules well separated by black ground. Hind wing, black, 
slightly duller and shading to more greyish on the inner fold. A postdiscal row of brighter 
blue spots extends from subcosta to above the anal angle, the line slightly bent outward at 5-4. 
The submarginal series of blue white-centred spots extends from the upper angle to anal angle 
where spots are double; marginal ochreous lunules, divided at mid point, may be distinct or 
faint; edge black, slightly dentate, tails black, rather short and sharply pointed, 4 and 3 mm 
long. Underside. Fore wing, ground colour brownish olive, with two irregular darker bands 
crossing the wing, separated by a black zigzag line on the proximal side of the discal zone. 
Cell with three black wavy bars, and stronger black lines sub-basal in ib and 20. In the post- 
discal line there is a series of slightly ochreous ocelli with dark centres, black in 2 and double in 
ib; the subcostal spot in this line is whitish. Margin with more ochreous lunules separated by 
dark ground. Hind wing, ground colour as fore wing, slightly more brownish at base, which 
has three olive-ochreous spots thinly outlined in black; discal line with interrupted ochre-olive 
marks outlined proximally in black, followed by less strong marks of the same colour in the 
postdiscal line. Submarginal row of whitish spots, black on distal edge complete, ending in the 
double marks at anal angle where the ground colour is olive. Margin with faint broken lunules; 
edge narrowly black. 

Female. Fore wing length 47-52 mm. Upperside. Fore wing, basal area brownish olive, 
shading to blackish in upper part of cell and on proximal side of white wing bar, which extends 
from costa at about end of cell to just short of the hind angle, the bar consisting of three elongate 
marks at end of cell followed by a more quadrate mark in 3, a larger more obliquely shaped 
mark in 2, below which are two smaller, more ochreous spots in ib in the hind angle. The distal 
portion of the wing is black with two angular white marks in the upper postdiscal line. Margin 
with two ochreous spots at hind angle in ib, edge immaculate or occasionally with minute 
whitish internervular spots. Hind wing, ground colour brownish olive with a strong black 
distal border. An occasional whitish ochre spot is present on the subcosta in the discal line 
and a series of ill-defined paler ochre-olive spots or lunules on the proximal side of the black 
border. Border with a series of submarginal blue white-centred ovoid marks extending from 
the upper angle to the hind angle where the spot is doubled. Margin with rather broken narrow 
golden ochre lunules, strongest above upper tail then fading out; edge bluntly dentate, tails 
7 mm and 4 mm. Underside. Fore wing, ground colour olive-greyish, more olive-brownish at 
base, wavy black lines narrowly outlined in white, present in the cell; bars at sub-bases ib-2 
stronger; black lines outlining the proximal border of the white bar strong. The white bar as 
upperside, the mark in ib contiguous with the postdiscal ochreous and black mark at tornus 
and space above, the rest of the spots in postdiscal line smaller, more greyish ochre; the two 
subapical spots whitish; the submarginal series of greyish lunules become progressively more 
faint to apex. Edge with very small whitish internervular marks. Hind wing, more or less 
as in the male, but ground colour browner olive. 

Descriptions taken from specimens from Ivory Coast and Ghana. 



REVISIONAL NOTES ON AFRICAN CHARAXES 237 

Range: The nominate race occurs in Sierra Leone, Guinea, Ivory Coast, Ghana 
and Fernando Po. 

Charaxes tiridates intermediate cline 

(Map 5) 

Charaxes tiridates ab. tristis Schultze, 19 14. 

Specimens from this area are generally larger, pattern of upperside very similar but blue 
spots in fore wing usually slightly more distinct; marginal lunules bolder. On the hind wing 
the blue spots are bolder, especially those of the postdiscal series; marginal golden lunules 
narrow or broken. Underside very similar to nominate race but basal black bars stronger. 

Range: Eastern Nigeria, Cameroun, Central African Republic, Congo (Brazzaville) , 
western Congo, Kasai and Katanga to Lake Mweru. Also N. Angola. 

Charaxes tiridates tiridatinus Rober 

(PI. 7, figs 51, 53, 55, 56, Map 5) 

Charaxes tiridates tiridatinus Rober, 1936 : 577, f. 3. 

Rober described tiridatinus as a 'form' or subspecies; the type was from Uganda. 
Since Uganda specimens are distinguishable from nominate tiridates of the Gold 
Coast, I accept this name for the Uganda subspecies. 

Male. Fore wing length 50-55 mm, majority 53 mm. Upperside. Fore wing, ground 
colour is a richer blue-black, slightly duller at base. The discal and postdiscal blue spots are 
larger, those of the discal line often extending to ib where the spot is in contact with the post- 
discal mark in the same area. The marginal golden lunules are larger. On the hind wing, the 
ground colour is blue-black, duller at base and shading to brownish on the inner fold. The post- 
discal blue spots are larger and there is often an additional spot at the subcosta in the discal 
line. The submarginal blue spots with white centres often arrow-shaped with an extension 
distally. The golden marginal lunules usually very strong and well marked; the edge with 
golden fringe indented by black dentate margin; tails short, upper 4-5 mm, lower 2-3 mm. 
Underside. Very similar to that of the nominate, the ground colour rather more golden 
brown; the sub-basal black bars less in evidence, but the overall pattern is the same and there 
is some variation in the olive-ochreous lunules on the distal half of the wings. 

Female. Very similar to the nominate, but generally larger, fore wing length 57 mm, the 
largest 59 mm, the smallest 55 mm. Upperside. Fore wing, general pattern is similar, the 
ground colour brownish olive, sometimes with a golden flush especially to the fore wings. The 
black shading on the proximal side of the white bar more in evidence especially in the upper 
part of the cell, the costa, however, olive-brown except opposite the bar where it is whitish. The 
bar is usually white, except at the hind angle, where the spots are ochreous. In some examples 
the bar is strongly yellow-ochre, the spot in 4 usually smaller than in the nominate race. In the 
black distal half of the wing, the two sub-apical spots are large, and there may be ochreous 
spots in the postdiscal line reaching to 2, in some specimens. The two ochre spots in ib at the 
tornus very distinct. On the hind wing the ochre costal spot is usually present, and in the 
postdiscal line the subcostal ochre-olive lunule is present, followed by a series of lunules on the 
inner side of the black border and reaching the anal angle, where they become obscured. The 
black border, widest at 6-7, tapers gradually, terminating at the anal angle. The series of 
blue spots with white centres complete, double at the anal angle. The marginal golden lunules 
usually very well marked, the spot at upper angle often large. The edge is narrowly golden 



238 



V. G. L. van SOMEREN 




-c -c -c -c 
<o u u u 



< 



REVISIONAL NOTES ON AFRICAN CHARAXES 239 

ochreous with slightly white fringe. Edge of wing bluntly dentate, tails slender, upper 6 mm, 
lower 5 mm, black with narrow white along lower edge. Underside. Fore wing, base ochreous 
olive, with strong black lines outlined in white; the black marks sub-basal in ib-2 strong. 
The distal portion of the wing darker, more brownish, setting off the white bar which is lined 
proximally in black. The postdiscal series of ochreous lunules (upper one whitish), together 
with the dark centres and paler outer border, form indistinct ocelli, becoming more distinct 
towards the hind angle where the central marks are black, the tornal double mark outlined 
distally with greyish lilac. Margin ochreous, well marked; edge with white fringe. Hind wing, 
ground colour and pattern as in the male ; the postdiscal series of olive-ochreous lunules may be 
strong; the submarginal spots enhanced by black distally, double at anal angle, moderately 
clear; the golden marginal line of lunules distinct; fringe white. 

Range: Western Uganda from Bwamba, Semliki Valley, Toro, to Kayonza in 
Kigezi; also on the west side of Lake Victoria to central and eastern Uganda and 
N.W. Kenya. It also occurs in northern Uganda at W. Madi on the Metu Hills and 
adjoining S. Sudan. In Tanzania, this race occurs in the Bukoba district to the 
eastern shores of Lake Tanganyika. 



Charaxes tiridates marginatus Rothschild & Jordan 

(PI. 8, figs 57, 58, Map 5) 

Charaxes tiridates marginatus Rothschild & Jordan, 1903 : 539. 

Male. Fore wing length 50 mm. Upperside. Fore wing disc deep blue-black, duller 
toward base, and black on outer border. Median line of blue spots well developed, as a rule, 
consisting of a streak and a quadrate mark at end of cell, two larger spots set out a little in 5-4, 
a trace of a spot in 2 approximating the postdiscal spot in the same area. Postdiscal series of 
blue spots complete, commencing at the subcosta and reaching ia, the line is outwardly curved 
in 5-6, and inwardly curved 4-3. The marginal golden orange marks, extending from the hind 
angle to the apex are strong, well developed, divided by the ends of the black veins. Hind 
wing, ground colour blue-black on disc, shading to dull greyish brown on the inner fold which is 
slightly ochre at the edge; border of wing black. Postdiscal series of blue spots complete, 
commencing at the subcosta and extending to above the anal angle, is slightly incurved at 6 and 
outwardly curved at 4. Submarginal blue spots with white centres complete and well marked, 
double at anal angle. Margin with strongly developed golden orange lunules, very slightly 
divided by black; edge black with buffish fringe, bluntly dentate; tails 5 and 3 mm long. Under- 
side. Ground colour slightly more greyish brown, the pattern not strong, but typical of tiridates. 
The marginal ochreous lunules of the fore wing and the border of hind wing pronounced. 

Described from specimens from the Leman River, S.W. Ethiopia (Hodson), 
kindty loaned to me by the Hope Dept., Oxford. Unfortunately no female is 
available. 

It will be noted, both from the description and the figures given, that marginatus 
comes very close to well marked specimens of the Uganda aggregate, and especially 
those from the northern districts of Uganda at Metu Hills, West Madi. 

Carpenter records tiridates from the Imatong Mts in southern Sudan, but does not 
comment on them. 

Range : The western and south-western areas of Ethiopia. 



2 4 o V. G. L. van SOMEREN 

SYSTEMATIC LIST 

Charaxes tiridates Cramer 

Charaxes tiridates tiridates Cramer, 1777. Type locality, Java (patria falsa). 

tiridates Cramer; Doubleday, 1844. Corrected type locality, 
Ashanti, Gold Coast. 

Synonym. Charaxes marica Fabricius, 1793. Locality 'Africa'. $. 

Charaxes marica Fabricius; Watkins, 1923. =tiridates Cramer <j>. 

Range : Sierra Leone, Guinea, Ivory Coast, Ghana and Fernando 
Po. 

Charaxes tiridates intermediate cline. 

Charaxes tiridates var. tristis Schultze, 1914. Type male, Bashe, Cameroun. 

Charaxes tiridates var. angusticaudatus Rober, 1956. Type male, Cameroun. 

Charaxes tiridates var. purpurina Rousseau-Decelle, 1938. 

Charaxes tiridates ab. subcaerulea Storace, 1948. Type locality, Congo, Etoumbe. 
Range: Eastern Nigeria, Cameroun, Central African Republic, 
Moyen Congo, west Congo, mid Congo River, Kasai, Katanga, 
Lake Mweru, Sudan, Nuba Mts. 

Charaxes tiridates tiridatinus Rober, 1936. Type male. Type locality, Uganda. 

Charaxes tiridates ab. conjuncta Storace, 1948. Type locality, Sesse Islands, 
Uganda. 

Range: Western Uganda, from Bwamba and Semliki Valley, Toro, 
to Kayonza in Kigezi ; also on the west side of Lake Victoria to 
central Uganda and N.W. Kenya; in Tanzania, this race occurs in 
the Bukoba district and the eastern shores of Lake Tanganyika. 
A smaller aggregate occurs in northern Uganda at West Madi on 
the Metu Hills and possibly on the adjoining hills in S. Sudan. 

Charaxes tiridates marginatus Rothschild, 1903. Type locality, Scheko, Ethiopia. 
Range: Western and south-western regions of Ethiopia. 



7. CHARAXES FUSCUS Plantrou and C. NUMENES Hewitson and 

its Subspecies 

Charaxes fuscus Plantrou 

(PI. 8, fig. 59, Map 6) 

Charaxes fuscus Plantrou, 1967 : 66. 

Rough translation by T. G. Howarth of the original description. 

Male. Body entirely brownish black. Fore wing length 40 mm, triangular in shape, the 
costa forming a very marked curve as in Charaxes protoclea Feisthamel and the distal margin is 
slightly concave. Hind wing, margin rounded and slightly crenate between each nervure with 
a tail present at veins 2 and 3, that at 2, 2 mm and that at 4, 3 mm in length. Upperside. 
Fore wing, ground colour velvety brownish black with a violet reflection in an oblique light 
with very few markings. A line of 7 small rather ill-defined postdiscal spots, paler than the 



REVISIONAL NOTES ON AFRICAN CHARAXES 241 

background, present in the interspaces. Hind wing, ground colour as fore wing with the same 
small postdiscal spots but these only present in spaces 5, 6 and 7. The outer margin is 
emphasized by a narrow reddish marginal band divided by the veins. The two anal spots are 
present in the form of two small bluish points but above these is a line of practically invisible 
subterminal points. Underside. Fore wing, ground colour brown similar to that of C. numenes 
Hewitson but a little more yellow. Four heavy and unequal black spots in the cell followed by 
two others at the middle of the costa. The third black cell spot from the base is adjacent to 
two black antemedian spots situated basad in spaces 1 and 2. A large tear-shaped black spot 
present in the postdiscal area of spaces 1, 2 and 3, then a subterminal line of pale spots bordered 
on both sides by dark spots, particularly accentuated in spaces 1 and 2. The inner margin is 
mostly darkened. Hind wing, two well marked black lines, both merging with the ground 
colour below the cell. An almost straight white median line bordered inwardly by a black line 
and outwardly by a blackish irregular line, then a complete series of warmer brown marks which 
form a projection in spaces 4 and 5. A subterminal line of six bluish streaks in line with the 
spots of the anal angle, finally a reddish outer margin. 

The holotype, a single male, was captured at Bangui, Central African Republic, 
in September 1966 by Monsieur R. P. Godart, and is in coll. Plantrou. 

The author places this newly described taxon immediately before C. numenes 
Hewitson. 

Charaxes numenes (Hewitson) 
(PI. 8, figs 60-63, Map 6) 

Kymphalis numenes Hewitson, 1865; t. 46. Type locality, Sierra Leone, types $ and $. 
Charaxes numenes Hewitson; Butler, 1865 : 630, n. 29. 

Nominate examples of Ch. numenes are small compared with their counterparts 
from Uganda and eastern Congo; they are also less blue-spotted, and differ in other 
ways. 

Male. Fore wing length 40-45 mm. Upperside. Fore wing, ground colour deep blue- 
black; the median or discal blue spots small, sometimes almost obsolete, a trace of a streak 
beyond cell with a distinct spot below, followed by two spots, upper one distinct, the other 
minute. The postdiscal row represented by two white spots, the upper one strong, the lower 
small or may be absent, these are followed by minute blue spots to ib, or these spots may be 
lacking. Margin of wing with distinct elongate ochreous marks, small and double in ib, larger 
in 2-3 then decreasing in size up to apex. Hind wing, ground colour blue-black shading to 
black on inner fold where the admargin is paler, but edge black. Blue postdiscal spots distinct 
from subcosta to just above the anal angle, the line of spots with a double curve; submarginal 
spots distinct, mostly white with slight blue surround, small and double in anal angle; margin 
with small divided triangular ochreous marks, more olive at anal angle; edge black with very 
narrow ochre fringe, very slightly dentate; tails very short, 3 and 1 mm. Underside. Fore 
wing ground colour greyish olive with ochre flush, the median area with a darker olive bar and a 
triangular patch beyond, base to costa; bold black spots and lines outlined in white in the cell, 
more rounded black marks sub-basal in ib-2; the two dark areas of the wing divided by an 
irregular black line strongly outlined distally in white; subapex with two whitish spots; border 
with indistinct ocelli with dark centres which gradually darken, the spot in 2 and double one 
in ib with black centres; basal area of ia greyish brown. Edge of wing with ill-defined ochre 
marks with black dots at vein ends. Hind wing, ground colour slightly darker than fore, basal 
half with a darker brown area divided at costal region by a pale line with two black spots 
proximally and a narrow black line distally which is edged with white proximally, the outer 
side of the dark patch accentuated sharply by an angled white line, narrowly black proximally. 



242 



V. G. L. van SOMEREN 




< 



REVISIONAL NOTES ON AFRICAN CHARAXES 243 

The postdiscal line with a series of ochre-brown lunules narrowly edged white on proximal 
side but shaded black distally. The submargin with a series of distinct white spots shaded lilac 
and black distally from 5 to anal angle where the spots are double. Border with broken tri- 
angular marks, ochre basally shading to white, olive at the anal angle; edge narrowly black with 
light brownish fringe. 

Female. Fore wing length 43-47 mm. Upperside. Fore wing, ground colour brownish 
olive in basal half, the distal half black, in between a bold white bar edged black proximally, 
strong at costa but decreasing to the hind angle; the white bar extending from the costa to just 
short of the hind angle, fairly even in width to 2, then with smaller spots which may be ochre- 
tinged. Sub-apex of wing with two large white spots, and occasionally with obscure spots in 
one to three spaces below. Margin with very slight indication of pale internervular spots. 
Hind wing, disc brownish olive usually stopping short at mid costa, where there is a pale 
subcostal rounded spot; beyond, a series of pale spots at the junction of the olive patch and the 
black border in the postdiscal line; the black border, widest at upper angle, tapers to above the 
anal angle, it carries a series of whitish lilac spots in the submarginal line. Marginal border 
very narrow, consisting of divided triangular ochre marks; tails stumpy, 5-2 mm. Underside. 
Pattern bold, as in the male. Fore wing basal half olive-drab, slightly more golden in the cell, 
which is crossed by bold lines and spots outlined in white; the sub-basal black marks in ib and 2 
bold, as are the black marks on proximal side of the white bar. On the distal side of the bar 
there is a dark triangular brown patch which extends up to the two subapical white spots; in 
the submargin there is a series of greyish ocelli with dark centres, which are black in 2 and double 
in ib at tornus. Border of wing more brownish, edge narrowly black with minute white fringe 
in interspaces. Hind wing very much as in the male; the postdiscal white and olive lunules, 
edged black, are stronger; the submarginal white and grey lunules well marked; margin with 
broken whitish triangles; edge greyish ochre. 

Range : Sierra Leone, Liberia, Ivory Coast and Ghana to western Nigeria. 



Charaxes numenes intermediate cline 
(PI. 8, figs 63, 64, Map 6) 

Charaxes numenes <$ i. laticatena Le Cerf, 1932 : 405, Belgian Congo, is a minor variation which 
occurs occasionally. 

Male. Fore wing length 45 mm. Upperside. Fore wing, ground colour as in the more 
western, nominate race. The blue spots in the median row rather obscured; the sub-apical 
spots smaller; the subcostal one distinct, but the remaining spots variable in number, hardly 
visible or absent. Marginal ochreous spots as in nominate or smaller. Hind wing ground 
colour as in nominate race; postdiscal blue spots smaller; admarginal dots more bluish; marginal 
line of small broken ochre triangles not reaching the upper angle; edge narrowly ochre-white, 
very bluntly dentate, tails stumpy 3 and 1 mm. Underside. As in nominate race. 

Female. Fore wing length 51 mm. Upperside. General pattern and colour as in nominate 
subspecies ; the white bar of fore wing less solid as the marks are more divided ; postdiscal spots as 
usual. Hind wing, colour and pattern as in more western examples. Underside. Ground 
colour as in nominate, but dark mid area in fore wing darker. In the specimen figured, the 
sub-basal black spots in ib-3 are conjoined to black marks on proximal edge of the fore wing 
white bar. 

Range: Cameroun, Central African Republic, Congo (Brazzaville), ? Kasai. 



244 V. G. L. van SOMEREN 

Charaxes numenes aequatorialis ssp. n. 

(PI. 9, figs 65, 66) 

Charaxes numenes ssp. et f. obsolescens Stoneham, 1931, Bull. Stoneham Museum, 7 : 1, Malawa 
Forest, Kakamega area, Kenya is of no importance. [Type examined]. 

Although there are no striking differences in the western and eastern aggregates 
the overall picture of the two, in series, above and below, is very noticeable. 

Male. Fore wing length 48-50 mm. Upperside. Fore wing, ground colour blue-black, 
slightly duller brownish at base. Median row of blue spots generally more distinct, often five 
in number, the subcostal one often whitish; the postdiscal blue spots more distinct; the sub- 
apical ones white and bluish white; the remaining ones more distinct than in nominate race; 
the marginal golden ochreous spots large and stronger. Hind wing basal area black more 
smoky on inner fold; border of wing blue-black; the postdiscal row of blue spots larger and 
brighter blue ; the submarginal white spots, blue distally, strongly marked ; the marginal golden 
ochreous triangles larger and less divided; edge golden, bluntly dentate, tails very stumpy, 
only 2 and 1 mm long. Underside. General pattern as in the nominate race, but often with 
the dark mid zone of the fore wing stronger, so also that of the disc of the hind wing ; the black 
marks in ib— 2 often connected by black bridges to the black marks in discal line. The black 
tornal spots strong, with golden ochreous surround; margin with stronger golden lunules, 
especially at hind angle. 

Female. Fore wing length 50-55 mm, larger than nominate females. Upperside. Fore 
wing, ground colour brownish olive with distinct rufous tinge shading to a narrow black line 
on the proximal side of the white bar ; the costal end of the bar more irregular than in nominate 
race, especially at 4 where the mark may be small, the whole bar more broken in outline, less 
solid, the marks often strongly yellowish tinged. Distal half of wing black with two well marked 
white spots in sub-apex, very rarely with trace of marks in 3 below. Margin with two ochreous 
spots in ib, and the slightest indication of marks in spaces above. Hind wing, ground colour 
as fore wing ; the black border widest at upper angle, may extend half way along the costa to the 
discal pale spot, tapering at the posterior end to above the anal angle ; on its proximal side is a 
row of paler postdiscal marks, most strongly developed at the subcostal end within the black 
ground of the border. Submarginal row of white, blue-pointed spots, double at anal angle, 
distinct ; admarginal row of golden triangles, usually larger than in nominate race ; edge ochreous 
with whitish elongate mark at upper angle. Underside. Fore wing, ground colour at the base 
of the wing as in nominate race but the dark brown area in the upper part of the disc wider and 
more angled distally, as a result, the white bar is narrowed; the black marks on the proximal 
side of the bar and those of ib and 2 either completely separated or slightly conjoined or joined, 
thus a variable character. Hind wing, the white edge to the discal brown band is stronger as a 
rule; the submarginal whitish lunules stronger; the amount of olive-ochre on the margin 
variable. 

Holotype male. Uganda: Kayonza, Kigezi, 6.1X.1952 (van Someren). 

Allotype female. Uganda: Kayonza Forest, Kigezi, v-vi.1957 (van Someren). 

Range : North-west Kenya and Uganda, extending into the eastern Congo along 
the Semliki Valley to Kivu; also found in the country south of Lake Victoria, 
extending south to east of Lake Tanganyika at Kigoma. 

Types in B.M.N.H. 

Charaxes numenes, local ecological form 

Representatives of numenes in the northern area of Uganda and northern Kenya 
and across the border into southern Sudan present an aggregate in which the males 



REVISIONAL NOTES ON AFRICAN CHARAXES 245 

are small, the wing length being 38-45 mm. The upperside colour is similar to the 
Uganda aggregate, but the ground colour is more brownish drab and the pattern 
not so bold. These may represent a cline toward the next subspecies. 



Charaxes numenes neumanni Rothschild 

(PI. 9, figs 67, 68, Map 6) 

Charaxes numenes neumanni Rothschild, 1902 : 597. 

Male. Fore wing length 45 mm. The characters given for the race are: marginal spots of 
both wings larger than in West African numenes; admarginal spots of hind wings also enlarged, 
confluent, yellow; white submarginal dots smaller than in numenes; discal spots a little more 
proximal. Underside, ground colour is more brownish, less olive drab, the darker bands on 
both wings dark brown, the white outlining very narrow. 

Described from a single male, this subspecies has now been recorded from Ethiopia 
by Ungemache at Youbdo; Hodson obtained it from the Ganji River and Haeto; 
Malcome Berkeley took it at Adola in south Ethiopia. 

SYSTEMATIC LIST 

Charaxes fuscus Plantrou 

Charaxes fuscus Plantrou, 1967. Type locality: Bangui, Central African Republic. 
Range: Only known from the locality of the unique type. 



C. numenes Hewitson 

Charaxes numenes (Hewitson), 1865. Types male and female. Type locality: 

Sierra Leone. 

Range: Occidental Africa: Sierra Leone, Liberia, Ivory Coast, 

Ghana, western Nigeria, Fernando Po. 
Charaxes numenes intermediate cline. 
Synonym: var. laticatena Le Cerf, 1932. Congo. 

Range: Cameroun, Central African Republic, Gabon, Congo 

(Brazzaville), N. Angola, Kasai, Katanga. 
Charaxes numenes aequatorialis ssp. n. 
Synonym: var. obsolescens Stoneham, 1931. Malawa, Kenya. 

Range: The eastern Congo from Semliki Valley to N.W. Kenya, 

including western Uganda to TransNzoia in Kenya; the area 

around Lake Victoria to eastern shores of Lake Tanganyika, in 

the north. 
Charaxes numenes aequatorialis ssp. Small aggregate. 

Range: Mt. Moroto, Turkana; Mt. Marsabit, Kenya. 
Charaxes numenes neumanni Rothschild, 1902. Type male. Type locality: Wori- 

Gamitscha; Kaffa, Ethiopia. 

Range : West and south Ethiopia. 



246 V. G. L. van SOMEREN 

8. CHARAXES BIPUNCTATUS Rothschild and its Subspecies 

Charaxes bipunctatus Rothschild 

(PI. 9, figs 69-72, Map 7) 

Charaxes bipunctatus bipunctatus Rothschild, 1894 : 536. 
Charaxes bipunctatus johnsoni Rousseau-Decelle, 1956. 

The type locality given by Rothschild, 1894, was 'West Africa, ? Gold Coast'; 
this was subsequently restricted by Butler (1896) to Accra, Gold Coast. In 1956 
Rousseau-Decelle described the race johnsoni, also from the Gold Coast, apparently 
thinking that specimens from further east, possibly Uganda, were nominotypical. 
In actual fact, the species bipunctatus is separable into two geographical races, with 
an intermediate cline between them. 

Nominate specimens from the Ivory Coast and Ghana are smaller than Uganda 
examples, and differ in other ways. 

Male. Fore wing length 43-44 mm. Upper side. Fore wing, ground colour blue-black. 
Blue spots in median or discal line variable, usually two streaks beyond the end of the cell, 
a slightly larger spot sub-basal in 3, a smaller spot in 2, not always visible. Subcostal spot in 
postdiscal line white, occasionally a blue spot below, and traces of spots in 2 and 3. Marginal 
ochreous lunules small, well separated by black ground. Hind wing ground colour blue-black, 
duller on inner fold, edge greyish, with slightly paler ochre tinge above anal angle. A subcostal 
blue spot at about mid point in the discal line ; postdiscal spots, two upper ones large, and may 
be followed by a trace of smaller spots in spaces below. Submarginal blue spots with white 
central dot complete, and double in the anal angle; marginal golden ochreous lunules narrow; 
edge black, slightly dentate, tails very stumpy. Underside. Fore wing, ground colour oliva- 
ceous drab at base, more olive greyish on the borders of wing. Cell with three wavy black 
lines, the central mark often divided, all white edged; the sub-bases of ib and 2 with larger 
black bars; the discal bar ochreous grey with a series of curved black lines on the proximal 
side; the submargin of the wing with rather obscure ocelli with darker centres more pronounced 
in ib and 2 where the centres are black, the marks outlined distally with bluish grey, the margin 
with rather obscure ochreous lunules most pronounced at hind margin. Hind wing, ground 
colour as fore, the base with wavy narrow black lines outlined white proximally, those of the 
discal line whitish distally, followed by obscure, slightly ochreous lunules with greyish and black 
proximally; submargin with small white dots, double at anal angle, which is distally shaded in 
black. Marginal border narrowly ochreous, edge black, more olivaceous at anal angle. 

Female. Fore wing length 45 mm. Upperside. Fore wing, base olivaceous brownish, 
shading to black in upper part of cell and proximal side of white bar, which crosses the wing 
from the costa to just short of the hind angle, where the spots are smaller and ochre-tinged. 
Distal portion of wing black with two white spots in subapex, occasionally a trace of a spot in 
4 below. Margin of wing with two ochreous spots at hind angle. Hind wing, ground colour 
lighter brownish olive; a whitish spot is present on subcosta at upper discal line, paler olive- 
ochre spots in upper part of postdiscal line becoming more obscure in lower half bordering on 
the black border, which is widest at upper angle and tapers toward the anal angle; submargin 
with a series of stellate white spots; margin with pale ochreous linules; edge black, slightly 
dentate, tails short and stumpy. Underside. Fore wing, ground colour and pattern as in the 
male but with the white bar, similar to above, but proximally outlined in black. Tornal marks 
black, but the rest of the ocelli above obscure. The two white sub-apical spots distinct. Hind 
wing more or less as in the male, but anal ocellus more distinct. 

Range: Nominate bipunctatus appears to be limited to the Ivory Coast and 
Ghana. I cannot trace any records from western Nigeria. 



REVISIONAL NOTES ON AFRICAN CHARAXES 



247 




< 



248 V. G. L. van SOMEREN 

Charaxes bipunctatus cline 

(Map 7) 

It is unfortunate that Rothschild gives such a wide distribution for the species 
in his monograph of the Charaxes (1900 : 390), and still puts the type-locality as 
'West Africa, ? Gold Coast', though accepting the specimens from Accra and Ashanti 
as nominotypical. Specimens from Cameroun and the Central African Republic, 
Gabon and possibly Moyen Congo appear to be intermediate in character between 
the nominate bipunctatus and those from Uganda and N.W. Kenya. It will be 
noted that the above description given by Rothschild is somewhat different from that 
of the original description (1894 : 536-537) based on the type. One can only 
assume that the latter was based on the augmented series then at Tring, which I 
suggest was a composite one. 

Male. Intermediate in size between the nominate race and that found in Uganda. Fore 
wing length 45 mm. Upperside. Fore wing, the blue spots in the median line are very obscure, 
sometimes almost absent; the subapical spot small; the marginal golden lunules stronger than 
in nominate. Hind wing, blue spots at upper angle strong or obscure; the submarginal small 
blue spots very faint and most marked at the upper angle; the marginal golden lunules strong. 
The characters of this cline thus embody those of the nominate and the Uganda race. 

Range: Cameroun, Central African Republic, Gabon, Moyen Congo and perhaps 
the Mid-Congo River region. 



Charaxes bipunctatus ugandensis ssp. n. 

(PI. 10, figs 73-75, Map 7) 

Male. Fore wing length 49-50 mm. Upperside. Fore wing, ground colour more purplish 
brown-black, with slight greenish sheen at base of wing. Discal or median line of blue spots 
usually small and limited to two, the uppermost beyond the cell usually absent, spots deeper 
blue, the spot in 3 is rarely present. Subapical spots, upper one large, distinct and white, the 
second spot small and obscured. Marginal golden lunules bold. Hind wing, ground colour 
as fore wing shading to brownish on the inner fold; subcostal blue spot in the discal line usually 
absent, but the upper spots in the discal line, one above the other, darker blue; very rarely a 
third spot. Submarginal spots blue or blue with white centres, strong in upper angle but tailing 
off towards the hind angle where the spots may be small and whitish ; marginal golden lunules 
strong and bold, very narrowly separated by black; edge black with ochreous fringe enhancing 
width of golden line; margin bluntly dentate, tails very stumpy. Underside. Ground colour 
browner; pattern similar but bolder except on the distal border where it is more obscured. 

Female. Fore wing length 50-54 mm, general colour and pattern similar to nominate female, 
but differing as follows: Upperside. Fore wing, the basal ground colour brownish olive, the 
amount of black shading toward the cell and proximal side of white bar less than in the nominate 
race ; bar similarly formed but third upper spot at base of 4 usually smaller and the marks more 
separated, those towards the hind angle more tinged with ochreous; the golden ochreous marginal 
spots larger and more distinct. Distal half of wing black, the two sub-apical spots larger, and 
there may be traces of spots, more ochreous in colour, in 4 and 5. Hind wing, ground colour 
as fore wing; a paler subcostal spot in discal line, then two light spots in upper postdiscal line 
with less conspicuous marks in spaces below; border of wing blackish, widest at upper angle 
where it may extend and encroach on costa, at lower end it tapers towards the anal angle; 
border with small whitish dots more in evidence at upper angle but less clear than in western 



REVISIONAL NOTES ON AFRICAN CHARAXES 249 

examples; border with wider golden ochreous marginal band, spot at upper angle more con- 
spicuous and rounded, the marks forming the band only narrowly separated by black veins; 
edge black with narrow ochreous fringe. Edge bluntly dentate, tails very stumpy. Underside. 
Fore wing, ground colour very like nominate race, slightly more brownish tinged, but the dark 
band through the disc stronger; wavy black bars in cell similar, but black marks sub-basal in 
ib-2 and proximal to the white bar larger; pattern on distal portion of the wing very similar. 
Hind wing ground colour as in nominate race, so also the general pattern, but less strong; black 
edge to postdiscal lunules less marked; submarginal whitish spots less in evidence, but these 
differences are slight. 

Holotype male. Uganda: Toro For. (J,), x.1949 (van Someren). 

Allotype female. Uganda: Toro, Kibali Forest, v-vi.1956 (van Someren). 

Within this general area, that is, in the Madi area of northern Uganda and neigh- 
bouring southern Sudan, we hnd a smaller aggregate, $ 45 mm, 9 47 mm. The 
males have a stronger greenish sheen at the base of the fore wing. The discal blue 
spots at the end of the cell more distinct ; the subapical spot strong ; golden borders 
of both wings strong. The females have slightly darker brown colour to base of 
fore wing and hind wing. The fore wing bar strongly tinged with ochreous. On 
the underside, the females exhibit a stronger pattern, the black marks sub-basal 
in 2 and 3 conjoined forming an almost solid black patch. This is probably merely 
an aberration. 

Range: Uganda and North-west Kenya, extending westward into the Semliki 
Valley and eastern Congo. The smaller aggregate is found in West Madi on the 
Metu Hills; and adjoining Southern Sudan. 

Types in B.M.N.H. 

SYSTEMATIC LIST 

Charaxes bipunctatus Rothschild 

Char axes bipunctatus bipunctatus Rothschild, 1894. Type locality: Gold Coast. 

Synonym. Charaxes bipunctatus johnsoni Rousseau-Decelle, 1956. 
Type locality: Gold Coast. 

Range: Limited to occidental Africa: Ivory Coast and Ghana, 
? western Nigeria. 

Charaxes bipunctatus intermediate cline to ugandensis. 

Range: Cameroun, Central African Republic, Gabon, Congo 
(Brazzaville) and perhaps the mid Congo River region. 

Charaxes bipunctatus ugandensis ssp. n. 

Range : Uganda and north-west Kenya, extending westward into 
Semliki Valley and eastern Congo. A smaller aggregate is found 
in West Madi on the Metu Hills and adjoining southern Sudan. 

9. The problem of CHARAXES MIXTUS Rothschild, 
CHARAXES BUBASTIS Schultze, and related species 

The males of both these species resemble somewhat the male of the Charaxes 
tiridates association, i.e. tiridates, bipunctatus, numenes, etc., whose females also 
c« 



250 V. G. L. van SOMEREN 

bear a resemblance to each other, having a brownish olive colour to the bases of 
fore and hind wing and having a conspicuous oblique white bar in the fore wings. 
The females of the mixtus group, on the other hand, are somewhat male-like, but 
duller and more brownish. 

It is of interest to note that at one time male mixtus was considered to be a 
variation of iiridates, and that bubastis was a hybrid between mixtus and smaragdalisl 

The male type of Charaxes mixtus came from Lokolele in the mid-Congo region; 
the type of male bubastis is from Bipindi in the Cameroun and there is a second 
example Irom the Njong River, also in Cameroun. 

In 1898 Rothschild described and figured what he took to be the female of Ch. 
mixtus. This specimen came from the Cameroun. It is totally different from any 
of the females of the tiridates association, and can be roughly described as having 
the appearance of a larger, duller, more brownish black form of the male. Later on, 
another female was taken at Kafakumba, Katanga, S. Congo. The association of 
these females with male mixtus has been generally accepted, but it has not been 
verified by breeding, nor by taking a pair 'in cop', so far as I can ascertain. 

Male mixtus is now well represented by specimens taken in the area from the 
Central African Republic, W. Congo, Kasai and Katanga. The female is still very 
rare, but two specimens have been kindly loaned to me, the type from the Berlin 
Museum, the other from the Congo Museum, Tervuren. They agree in all respects. 



Charaxes mixtus Rothschild 

(PI. 10, figs 76-79, Map 7) 

Charaxes mixtus Rothschild, 1894 : 554, t. 12, fig. 8. 

Charaxes mixtus Rothschild, 1898 : VI. Female, type locality: Cameroun. 

Male. Fore wing length 48-50 mm. Upperside. Fore wing, ground colour blue-black, 
duller black at the base. Discal or median row of blue spots rather variable in number, some- 
times three, sometimes four to five in a curve; the upper mark may be a mere blue streak in the 
subcosta, followed by a larger spot at the end of the cell and more conspicuous spots in 3-2 and 
the trace of a spot in the upper part of ib. The postdiscal row of spots consists of two well- 
marked white spots in subapex, followed by a trace of spots, blue in colour, in spaces below, 
the marks in ib and especially that in ia, bolder. The margin of the wing with whitish marks 
double in ib, strong in 2, then more diffuse in spaces up to the apex. Hind wing, ground colour 
blue-black, duller on the inner fold which is smoky grey-black. On the postdiscal line there are 
conspicuous blue spots in a row from subcosta to above the anal angle, the spot in 6 set in so that 
the line is here kinked or angled proximad. The submargin has a row of bluish white spots 
or stronger blue with white centres. Admargin with a series of bluish white lunules, sometimes 
white at upper angle; edge with whitish internervular fringe; margin slightly dentate. Tails 
thin and pointed, 5-3 mm long. Underside. Both wings drab olive-grey, slightly darker in 
disc of fore wing. Fore wing, cell and sub-bases ib-2 crossed by black lines slightly edged with 
white; discal line of paler angles outlined proximally in black not very strong; postdiscal series 
of obscure ocelli become strongly marked at tornus and space above; subapical spots whitish; 
margin with olive-ochreous lunules. Hind wing with thin black marks in sub-base; disc with 
paler spots finely edged with black proximally, followed by a postdiscal series, kinked at 5, 
extending from 6 to anal angle and crossing the inner fold ; submarginal series of whitish spots, 
blackish distally, lie in contact with the admarginal series of olive-ochre lunules; margin 
brownish black with paler fringe. 



REVISIONAL NOTES ON AFRICAN CHARAXES 251 

Female. Fore wing length 54-56 mm, thus larger than the male. Upperside. Ground 
colour of both wings rather browner especially at the bases. Fore wing, median blue spots 
larger but duller, that at end of cell obscured. Postdiscal spots obscured, except the two at the 
hind angle which are dull bluish while the two subapical are clear and white, the upper one 
somewhat linear. Marginal spots rather obscured, slightly ochreous in colour, double in ib. 
Hind wing, pattern of blue spots as in the male, but postdiscal spots almost obscured except 
for that in 6 but even this is dull. Submarginal bluish spots complete but dull; marginal 
lunules well developed but dull olive-ochreous. Tails, rather thin and pointed, upper 7 mm, 
lower 4 mm. Underside. Fore wing, ground colour brownish olive, paler at the base and with a 
darker zone in the discal area; black wavy lines cross the cell, that at end of cell adjacent to the 
series of black lunules distally shaded with greyish, in the median line; the postdiscal series of 
strongly ochreous spots in ib and 2 become increasingly obscure but more whitish in the sub- 
apex; the tornal black marks, double in ib are strong. Margin with ochreous marks decreasing 
in size from ib to apex. Hind wing, ground colour as in fore wing; basal black lines thin but 
distinct corresponding to those in the male. The postdiscal series of paler spots in the form of 
lunules, complete but weak, that at anal angle stronger. Submarginal spots complete, dull, 
sometimes with black dot distally, double at anal angle on olive ground. Admargin with 
ochreous lunules, edge darker brownish. 

Variation: The amount of variation in the male is not, as a rule, very great as 
can be seen from the figures on Plate 10, with the exception of fig. 77, which depicts 
a specimen with large and extended blue spots in the postdiscal zone of the hind 
wing. On the other hand the blue spots on the upperside of the type (PI. 10, fig. 76) 
especially in the hind wing, are obscured. 

There is nothing on record regarding food plant or early stages. 

Range: Katanga, S. Congo; W. Congo; Central African Republic; Cameroun. 



Charaxes bubastis Schultze 
(PI. 10, fig. 80, Map 7) 
Charaxes bubastis Schultze, 1917 : no, t. 13. 

As indicated in the introductory note, there has been some speculation as to the 
correct status of this Charaxes. The suggestion that it is a hybrid can bo discounted 
for such are extremely rare in nature. Moreover, the insect is known from at least 
four specimens taken in different localities, at different times. 

I have before me two specimens, one compared with the type and another perfect 
example ; they agree in all respects. 

Male. Fore wing length 51-52 mm. Upperside. Fore wing, ground colour blue-black 
with a slight purplish tinge, more brownish at the base. Median or discal row of blue spots 
larger than in mixtus, more purplish blue, consisting of a streak beyond the end of the cell, 
followed by a smaller more rounded spot sub-base in 3, followed by a larger spot directly below 
in 2, the double spots in ib (vestigial in one specimen). In the postdiscal row, the subapical 
spots are bold and white, while the lower spots are obscured, though more apparent in ib and 
ia. Margin with well marked ochreous spots well separated by dark ground, double in ib, and 
extending up to the apex; these spots more defined than in mixtus. Hind wing, ground colour 
purplish blue-black shading to more greyish on the inner fold. Disc of wing in one specimen 
with a slight greenish bloom; there is also a subcostal blue spot. Postdiscal row of spots may 
be complete or lacking the upper spot in subcosta, the remainder blue with slight purplish tinge, 



252 V. G. L. van SOMEREN 

the upper three spots less angled than in mixtus and nearer the submarginal row of whitish blue 
spots, which in turn are less approximated to the marginal row of blue lunules, which are whitish 
at the upper angle ; edge slightly serrate ; tails more robust and shorter, 4-3 mm long. Under- 
side. Fore wing, ground colour olive-drab, more olive-ochreous at basal angle, discal area 
rather darker. Cell crossed by the usual black lines, narrowly outlined in white; a median 
series of light lunules accentuated in black proximally present; stronger black lines sub-basad 
in ib and 2. Submarginal ocelli with black centres present in ib and 2, the rest tending to fade 
out; subapical white spots strong; margin with olive-ochre spots not as defined as on upper side. 
Hind wing, ground colour as fore wing; very thin black in basal area; discal and postdiscal 
paler spots distinct, those in the lower portion of the line with dark shading distally; sub- 
marginal row of spots as above but not so defined; marginal lunules greyish buff; edge brown 
not sharply serrate but more dentate. Anal angle golden olive with double black dots. 
The female is not known. 

Thus there is some general resemblance to Charaxes mixtus ; but since the differ- 
ences between the two are obvious, I consider bubastis a distinct species. 

Range: Known only from Cameroun, specimens having been taken at Bipindi 
(type-locality) and at the Njong River. 



Charaxes albimaculatus sp. n. 

(PI. 11, figs 81, 82) 

Amongst the material ot 'mixtus kindly loaned to me by the British Museum 
(Nat. Hist.) is a male specimen which had been tentatively placed as a variation of 
mixtus Rothschild. It, however, exhibits some outstanding characters which 
suggest that it belongs to a distinct undescribed species. This specimen was taken 
at Stanleyville in northern Congo. 

Until recently no lemale could be associated with this unique male but, as a 
result of exchange of photographs and specimens with Monsieur Plantrou of Paris, 
it now appears that he has in his collection a female which belongs to this species. 
The specimen was obtained at the recent sale of the Le Moult collection in Paris. 
This insect has now been forwarded to me, and it is without doubt a female of the 
male I am now describing. It possesses all the essential characters of the male, 
including the conspicuous submarginal white spots in the hind wing above and the 
almost uniform colour of the hind wing below, thus differing considerably from the 
female of mixtus, to which it had been placed. 

Male. It is considerably smaller in size than mixtus Rothschild, fore wing length 42 mm; 
the hind wings more pyriform due to the more pointed anal angle; the antennae are shorter. 
Upperside. Forewing, ground colour is a deep blue-black, slightly duller blackish at the base. 
The blue spots in the median and postdiscal line are small and punctiform, the two upper spots 
in the median line larger than the rest ; the ochreous spots on the margin are smaller but more 
distinct. The hind wing is blue-black, shading to brownish on the inner fold, immaculate in the 
disc and postdiscal area, but the submargin carries a row of conspicuous white spots, double at 
anal angle; this character is outstanding. There is no marginal border, the wing being black 
to the edge which is bluntly serrate. Underside. Fore wing, strongly patterned, the ground 
colour a warm olive brownish, paler, more ochreous at the base where the black lines and spots 
are bold, that crossing the end of the cell continuous with the black marks in sub-bases of ib 
and 2. The mid zone of the wing is obliquely crossed by black lines distally shaded in ochreous 



REVISIONAL NOTES ON AFRICAN CHARAXES 253 

to whitish, terminating in a large black mark in ib, which is contiguous with the darker zone 
separating the oblique bar from the rather obscure ocelli of the postdiscal line, the subcostal 
subapical mark whitish; the ocelli in ib and 2 with black centres; margin with ill-defined ochreous 
lunules, double at tornus. Hind wing, ground colour almost uniform brownish olive with one 
faint subcostal spot in median line; black line in sub-base very thin, median area and postdiscal 
zone without marks, but submargin with white spots as upperside; margin slightly darker 
especially in mid area, but there is no marginal border. Anal angle with lilac and double black 
dots. 

Female. Fore wing length 54 mm. Upperside. Fore wing, ground colour purplish brownish 
black at base of wing but darker in distal half; purplish blue spots in the discal line from end 
of cell to ib where the marks are faint; two white spots in the subapex, the upper one rectangu- 
lar, the lower smaller and rounded; margin with faintly indicated ochre-greyish spots; edge 
faintly whitish. Hind wing, basal area as fore wing, shading to greyish brown on the inner fold ; 
distal portion of wing darker brownish black with indication of three paler purplish brown spots 
in postdiscal line in 6-7, these spots placed more toward the series of conspicuous white sub- 
marginal, somewhat angled, spots placed as in the male. There are no marginal lunules; 
extreme edge faintly whitish. Tails short, 4 and 3 mm. Underside. Similar in all respects 
to that of the male. 

Holotype male. N. Congo: Stanleyville, iii.1924 (Ertl collection, ex Joicey 
Bequest, B.M.(N.H.). 

Allotype female. S. Congo: Katanga, Kafakumba (Overlaet Collection, ex Le 
Moult) in Coll. Plantrou, Paris. 



Charaxes barnesi Joicey & Talbot 

(PI. 11, figs 83, 84) 

Charaxes barnesi Joicey & Talbot, 1927 : 14. 

No actual specimens are available to me for description but, through the kindness 
of the British Museum (Nat. Hist.), photographs of the types are here reproduced. 
For the brief description I have had to draw on the original one given in the publica- 
tion cited above. 

Male. Fore wing length 48 mm. Upperside. Ground colour of both wings deep blue, 
more intense over the proximal half of both wings. Fore wing, the median row of blue spots 
consists of three streaks in subcostal area in 4-6, followed by more quadrate marks set out a little 
in 3-2, followed by a smaller spot in upper part of ib approximating toward the postdiscal 
spot in the same area; post discal spots complete, commencing with two large white spots in 
subapex, followed by blue marks of increasing size and extending to the hind margin where the 
mark is a streak. Margin with white linear marks, double in ib, decreasing in size up to apex. 
Hind wing, disc immaculate, but postdiscal zone with a series of large blue soots from subcosta 
to above anal angle, the spot in 6 set in so that the line is here kinked, the jpots in 3 set in a 
little so that the line has a double curve; submargin with a row of blue-white centre spots; 
marginal border with whitish lunules; edge black, very slightly dentate, tails comparatively 
short and thick, upper 5 mm, lower 2 mm. Underside. Fore wing, ground colour olive-drab; 
black bars narrowly outlined in white in the cell; thicker black bars sub-basal in ib-2, these 
marks set in more basad than usual. Disc of wing crossed by a paler zone in the median line, 
corresponding to the blue of upperside, but extending to and tapering in ib. Two white 
subapical spots prominent ; postdiscal area with paler lunules forming the inner side of the row 
of obscure ocelli with darker centres, more distinct in ib where the centre is black. Hind 



254 v - G. L. van SOMEREN 

wing, ground colour as fore wing; base with small light marks outlined in black; discal zone with 
thin black line shaded whitish distally, crossing the inner fold ; postdiscal row of light yellowish 
marks corresponding to blue marks above; submarginal series of whitish spots double at anal 
angle, clear; marginal border pale ochre. 

Female. Upperside. Joicey and Talbot compare this with Ch. cithaeron as having the same 
purplish brown ground colour. The white band of the fore wing commencing at the subcosta 
as a streak, widens to 2, then abruptly narrows in ib, the double spots being set towards the 
end of the mark above. The margin has two pale spots in ib. Hind wing with the discal 
border blackish, defined on the inner border by a series of purplish blue spots, large and distinct 
at subcosta and decreasing in size and distinctness to above anal angle. There is a large well 
defined bluish spot in the subcosta at upper median line. A complete row of blue white-centred 
present in the submargin ; the margin with distinct lunules, white at upper angle, then shaded 
with purplish toward anal angle. Tails thick, relatively short, upper 6 mm, lower 3 mm. 
Underside. Fore wing, ground colour as in the male; the white bar formed exactly as above; 
the two subapical white spots strongly represented; the obscure submarginal ocelli as in the 
male; margin without any pale marks. Hind wing, ground colour as in the male, the pale 
mark at subcosta in the discal line followed by a zigzag series of less strong marks to above anal 
angle where the spot on inner edge of fold is whitish ; postdiscal series of paler spots arranged in 
same way as marks above ; submarginal row of whitish spots, double at anal angle fairly distinct ; 
marginal lunules whitish at upper angle are shaded with purplish in region of tails and anal 
angle. 

Range : Known only from the island of Principe in Gulf of Guinea. 



SYSTEMATIC LIST 

Charaxes mixtus Rothschild 

Char axes mixtus Rothschild, 1894. Type locality: Lokolele, mid Congo River (<J), 
Cameroun ($). 

Range: S. Congo, Katanga, W. Congo, Central African Republic 
(Moyen Congo), Cameroun. 

Charaxes bubastis Schultze 

Charaxes bubastis Schultze, 1917. Type locality: Bipindi, Cameroun. 

Range: Only recorded from Cameroun at Bipindi and Nijong 
River. 

Charaxes albimaculatus sp. n. 

Charaxes albimaculatus sp. n. Type locality: N. Congo, Stanleyville (<$), S. Congo, 
Kafakumba, Katanga. 
Range: N. and S. Congo. 

Charaxes barnesi Joicey & Talbot 

Charaxes barnesi Joicey & Talbot, 1927. Type locality: Principe I. 

Range : Confined to Principe Island in the Gulf of Guinea. 



REVISIONAL NOTES ON AFRICAN CHARAXES 255 

10. Further notes on CHARAXES MANICA Trimen and 
C. MCCLEERY I sp. n. 

Charaxes tnanica Trimen 

After my previous notes on this species (1966 : 86) had gone to press, Monsieur 
Jacques Plantrou of Paris received, during the latter part of 1966, some interesting 
specimens of a 'Black' Charaxes of the 'Etheocles Complex taken in the Brazzaville 
area of the Congo by Monsieur Auberger. Monsieur Plantrou rightly placed them 
to the species manica, but noted that they differed considerably from the nominate 
race. The specimens were in due course submitted to me for an opinion. There 
is no doubt that they represent a good subspecies from an area in which manica 
has not hitherto been recorded. 



Charaxes manica subrubidus ssp. n. 

(Pis 11, 12, figs 85-92) 

Male. Fore wing length 36 mm. Upperside. Fore wing, shape and ground colour similar 
to the nominate race. The subapical spots slightly larger and whiter, the blue spots beyond 
the end of the cell more obscured, that at upper part of end of cell barely indicated. Hind 
wing submarginal bluish spots as in nominate race, but the marginal border slightly broader 
and brighter red above upper tail but mixed with golden olive from lower tail to anal angle; 
edge black with trace of a fine white fringe. Tails as in the nominate race. Underside. Fore 
wing, ground colour and pattern as in nominate but slightly more rufous at base; the postdiscal 
lunules larger, more distinct and reddish. Hind wing, ground colour more reddish toward 
base; pattern similar, but postdiscal lunules strongly reddish and mixed with orange at the 
anal angle. Marginal border wider and brighter red to upper tail, then mixed with orange to 
anal angle. 

Holotype male. Congo: Brazzaville, October 1966 (coll. J. Plantrou). 

The female forms taken so far are also distinctive, though some of them have 
their counterpart in the females of manica manica. 



$ form atribasis forma n. 
(PI. ii, figs 87, 88) 

Fore wing length 40 mm. Upperside, nearest to the form manica of the nominate race but 
differing as follows: Fore wing, base blackish, or blue-black in side light as far as the upper 
part of the white oblique bar, the pale blue being limited to the lower proximal half of the bar 
in ia-ib and very slightly in 2. The white bar is widest at the costa where there is an extension 
into the sub-bases of 6-7, the bar tapering slightly to 2, then represented in ib by a smaller 
spot in upper part and a few white scales in the lower which is mainly blue; the distal half of the 
wing is black; the subapical whitish spots large; edge of wing with very narrow white inter- 
neural fringe. Hind wing, basal area black with bluish bloom in side light, shading to more 
greyish on the inner fold; disc of wing with a broad pale blue discal patch, commencing at the 
costa and widening in 5-4, then tapering to above anal angle; border of wing black, widest at the 
upper angle with a complete row of rather conspicuous lilac spots with white centres in the 



256 V. G. L. van SOMEREN 

submargin. Marginal border brick-red above upper tail then mixed with olive toward the hind 
angle; and edge black; tails fairly long and thin, 6 and 5 mm long. Underside. Fore wing, 
rather paler than in nominate, the satiny greyish brown area in subapex more distinct. The 
extension of the white bar along the subcosta more strongly marked than above; the black 
tornal marks and the black marks on proximal side of white area in ib, strong. Hind wing, 
ground colour not so dark, but the postdiscal row of reddish lunules conspicuous as are the white 
lunules in the submarginal line, those toward the anal angle distally accentuated with black. 

Holotype female. Congo: Brazzaville, 19. x.1966 (coll. J. Plantrou). 



$ form aubergeri forma n. 

(PI. 12, figs 89, 90) 

Fore wing length 40 mm. Upperside. Fore wing, pattern corresponding more or less to the 
form chintechi of the nominate race, the orange spots of the discal and postdiscal row bolder, 
with an extension into the cell at 4, but these spots are on a greenish black ground, those of the 
discal line suffused over with a beautiful iridescent greenish blue which replaces the white of the 
nominate form, and extending basad onto the black base of the wing. Margin of wing with 
obscure ochreous spots, white and double in ib. Hind wing, discal and postdiscal area with a 
large patch of iridescent bluish green, slightly paler in the disc and slightly shaded orange 
toward the costa where there is a distinct blackish oval mark; the base of the wing dark but 
with a blue sheen. Border of wing strongly black, carrying a complete row of conspicuous 
bluish white linear marks; margin with reddish lunules outlined in white mixed with olive 
toward the anal angle. Tails long, thin, upper 6 mm, lower 5 mm. Underside. Fore wing, 
ground colour strongly rufous, the discal and postdiscal spots on upperside are here dull orange, 
the marks in ib coalescent; tornal black mark strong, with two whitish dots distally on margin. 
Hind wing, ground colour rufous brown with a paler disco-postdiscal bar crossing the wing 
accentuated proximally by a fine black line and distally by more reddish contiguous lunules in 
the postdiscal line. Submarginal white linear marks strong; marginal red border rather narrow- 
outlined greyish proximally shading to olive at anal angle. 

Holotype female. Congo: Brazzaville, 25.V.1967 (coll. J. Plantrou). 
This distinct form is name after its discoverer. 



$ form pseudosmaragdalis van Someren & Jackson comb. n. 
(PI. 12, figs 91, 92) 

Charaxes cedreatis vetula, ? f. pseudosmaragdalis van Someren & Jackson, 1957 : 89. 
Charaxes cedreatis, ? f. pseudosmaragdalis van Someren & Jackson; van Someren, 1969 : 89, 
pi. 3, figs 18, 19 (type). 

This distinctive form was erroneously assigned to Charaxes cedreatis, largely on 
account of its very conspicuous submarginal linear marks above and below on the 
hind wing. Moreover, Ch. manica had not been recorded further west than Katanga. 
I am glad to have the opportunity of correcting the error. 

Fore wing length 40 mm. Upperside. Fore wing, ground colour, distal portion black, 
basal area black at end of cell, but strongly suffused with greenish blue to the base, as in the 
other females described. The wing is crossed by two rows of spots, discal and postdiscal, 
exactly as in f. aubergeri, the marks coalescing at the hind margin; the discal spots are bright 



REVISIONAL NOTES ON AFRICAN CHARAXES 257 

blue, as are the postdiscal, except for the two subapical ones which are white. The margin 
of the wing has a series of obscure pale spots as in aubergeri. Hind wing, basal area is blackish 
with the same strong greenish blue iridescence in side light; the inner fold is greyish. The disc 
of the wing has a large iridescent blue patch as in the other forms, the outer border sharply 
defined from the black border, which carries distinct bluish white linear marks, double at the 
anal angle; the marginal border is reddish above the upper tail, outlined in whitish and mixed 
with olive at the anal angle. Tails as in other forms. Underside. Fore wing, ground colour 
of one of the specimens taken at Brazzaville is more rufous than that of the type, but the satiny 
bars and the black marks at the hind angle are similar. The same remarks apply in regard to 
the hind wing, but unfortunately the distal portions of both hind wings are missing in the 
Brazzaville specimen submitted. 

Range: So far as is known, this new form of manica occurs in the region of Brazza- 
ville and Leopoldville, and greatly extends the range of the species to the west. 

Note by Monsieur Jacques Plantrou. 

'This form seems to be especially rare, and there is too little material to form an 
idea as to the relative proportions of the various female forms. As far as we know, 
only two males, three female atribasis, two aubergeri and three pseiidosmaragdalis 
have been taken in spite of intensive trapping by at least three collectors during a 
period of three years.' 



Charaxes mccleeryi sp. n. 

(PI. 12, figs 93, 94) 

Charaxes sp. n.? van Someren, 1969 : 163, No. 26. 

This species belongs to that highly complex association which includes manica, 
alpinus, etheocles, ethalion, etc., whose males are often confusingly alike and whose 
females are very variable. After a thorough examination of its genitalia along 
with those of allied species, one is forced to regard mccleeryi as a distinct species. 
Moreover its distribution coincides and is overlapped by others of the group. For 
comparative illustrations of genitalia, vide van Someren, 1969 : 163, No. 26. 

Male. Including the type, fore wing length 34-36 mm; apex bluntly pointed; outer margin 
slightly incised. Upperside. Fore wing, black, with very slight greenish tinge at base, immacu- 
late, except for two faint blue spots in the sub-apex, sometimes only one. Extreme edge of 
wing with very narrow white fringe in interspaces. Hind wing, black, slightly duller on inner 
fold; minute white dots on submargin, in region of tails, and double mauvish spot in anal 
angle; border of wing maroon above tails, shading to olive-green from upper tail to anal angle. 
Margin of wing very slightly serrate, tails at veins 4 and 2, 5 and 6 mm long, thin, black in 
colour with olive centre line. Underside. Fore wing, ground colour warm greyish brown with 
ferruginous bloom; satiny bars in disco-postdiscal line divided at costa by a dark quadrate mark. 
Three ovoid black spots in fore wing cell narrowly outlined in white, fine black lines beyond, 
those in sub-bases 6-2 in discal line, black. Submarginal zone with obscured dark spots, but 
double and very distinct in ib at the hind angle, the black spots shaded with greyish proximally 
and outlined in black distally. Hind wing, ground colour as fore wing, the dark discal bar 
edged in black; postdiscal zone with maroon lunules edged with olive proximally and lined in 
black, the olive more apparent above the anal angle. Margin of wing maroon to upper tail, then 
olive, edge proximally in black, double spot in anal angle; edge black with narrow white fringe. 
There is a pale mark on the inner fold above the anal angle. 



258 V. G. L. van SOMEREN 

Female. Those captured to date bear a strong resemblance to the white-barred form of 
etheocles but with smaller discal-postdiscal white spots in fore wing. Fore wing length 
42 mm. Upperside. Fore wing, shape similar to the male but outer margin less incised. Ground 
colour brownish black, with slight greenish tinge at base. Spots in discal line as follows: two 
subcostal spots in sub-bases 6-5, upper larger, followed by spots in sub-base 3 and base of 4, 
the latter small and set in towards the end of the cell, the spot in 2 quadrate, than in ib larger, 
with incised inner edge, that in ia an elongate streak, the upper spots off-white, the lower 
white. The postdiscal row of spots are smaller and well separated, three in a row in sub-apex, 
spot in 4 set in, followed by an angular mark in 3 and a faint mark in 2. The two rows of spots 
are well separated. Hind wing, ground colour black in basal area and border, greyer on the 
inner fold. The disc of the wing with a white bar, 6 mm wide at costa then tapering but lower 
borders strongly bluish, especially on the proximal side. Border of wing with mauve lunules 
with white centres, ending in a double mauve spot in the anal angle ; margin with a maroon line 
shading to olive and expanding at anal angle. Tails long and thin, upper 7 mm, lower 8 mm. 
Underside. Fore wing, ground colour earthy brownish grey with rufous bloom; darker bar in 
the discal-postdiscal zone separating the two rows of whitish spots, those in the outer row 
bufhsh, both arranged as upperside. Border of wing with hardly any indication of dark sub- 
marginal spots except in ib, where the double black spots are large, conspicuous and bordered 
with whitish, forming a strong 'eye-spot'. Hind wing ground colour as fore; basal dark lines 
faint; discal white bar narrower than above and extending to just beyond the end of the cell. 
Postdiscal maroon lunules, edged olive and black proximally, fairly distinct, shading to olive 
above anal angle, with a pale bufhsh mark on inner fold just above. Marginal white lunules 
very distinct, bordered by equally distinct reddish lunules to upper tail then shading to olive at 
anal angle, with some black between the rows, especially in the region of the tails, ending in 
the double black spot in the anal angle. 

Some females are more flushed with rufous over the entire underside. 

Holotype male. Tanzania: Bunduki Hill, Uluguru Mts, 6000 feet. vii.T966 
(C. H. McCleery). To be deposited in the British Museum (Nat. Hist.). 

Allotype female. Same data. 

Paratype males (2), Uluguru Mts, February 1967 (/. Kleilland). 

Range: At present known only from the Uluguru Mts in Tanzania. 



SYSTEMATIC LIST 

Charaxes manica Trimen 

Charaxes manica subrubidus ssp. n. Type locality: Congo, Brazzaville. 
Range: Congo, Brazzaville, Leopoldville. 

Charaxes manica subrubidus $ f . atribasis forma n. Type locality : Congo, Brazzaville. 

Charaxes manica subrubidus $ f. aubergeri forma n. Type locality: Congo, Brazza- 
ville. 

Charaxes manica subrubidus $ f. pseudosmaragdalis van Someren & Jackson 1957, 
comb. n. Type locality: Congo, Leopoldville. 



Charaxes mccleeryi sp. n. 

Charaxes mccleeryi sp. n. Type-locality: Tanzania, Uluguru Mts. 
Range : Only known from the type locality. 



REVISIONAL NOTES ON AFRICAN CHARAXES 259 

11. Further notes on the CHARAXES XIPHARES Complex 
and a Description of a New Subspecies 
Charaxes xiphares woodi van Someren 
Charaxas xiphares woodi van Someren, 1964 : 195. 

When I described this subspecies of Charaxes xiphares, it was known from only 
two males. Intensive search has been made for the female in the two localities 
from whence the males were taken, but it was not until April 1966 that an almost 
perfect female was secured by Mr Peter Martin on Soche Mt., 5000 feet, Malawi, 
in almost the same spot where he had taken a male earlier in the same month. He 
took a second female at the Mlosa Stream in August 1967. 

Between these dates, two more females and a male were secured by Dr C. H. 
McCleery in the Nchisi Forest, Central Province, Malawi between January and 
March 1967, thus extending the range of this subspecies considerably to the north. 

The males agree in all essential respects with the holotype and paratype described 
in 1964, the only difference is in the width of the orange border of the hind wing, but 
this is shown to be variable, in specimens of both sexes, from the same locality. 

Female. Fore wing length 54-55 mm (neallotype 55 mm, a small specimen 50 mm). Shape 
very similar to that of the male, but slightly less incurved on the margin at 3-4. Upperside. 
Fore wing, ground colour black, crossed by a somewhat interrupted discal white bar consisting 
of a thin white area on the costa, two elongate and a triangular mark at bases of 6, 5 and 4, 
divided by black veins, a more obliquely quadrate mark in 3 set out a little, forming an angle 
with marks above, a long somewhat ovoid mark in 2 with a small rounded spots at its distal end 
in upper part of 2b and a more triangular mark in lower portion, followed by an elongate mark 
in ia extending proximad; the three lower marks in contact with the rather obscured orange- 
ochreous, rounded spots of the postdiscal line in these areas, the rest of the spots in the post- 
discal line free, forming a gentle curve with larger more whitish spots in sub-apex 6-7. Margin 
with golden ochreous spots, double at the hind angle, decreasing in size up to apex. In some 
specimens, the marginal spots, other than those at the hind angle, missing or only faintly indi- 
cated. Hind wing, ground colour black at base shading to ashy grey on inner fold; outer border 
broadly black; disc of wing with a large bluish patch, ill-defined on its lower half, becoming 
whitish and more defined and restricted in 4-6 to subcosta, the inner border encroached on by 
the black ground. On the distal side of the patch there are three ill-defined ochreous spots in the 
postdiscal line. Submargin with a complete row of somewhat triangular bluish spots. Marginl 
with strong line of golden ochreous lunules, slightly separated by black veins, olive green at anae 
angle; edge black with narrow white fringe. Tails short and stumpy. Underside. For 
wing, ground colour olive-greyish, slightly paler at the base; cell with four black lines, narrowly 
outlined in bluish white, sub-base of 2 with a short black bar, and traces of lines in sub-base ib. 
The discal area of the wing darker, with the white bar as above, except that the white marks in 
ib are smaller; and more defined. The postdiscal row of ochreous spots, more in the form of 
lunules from ib-3, accentuate the proximal edge of the conspicuous black marks at the hind 
angle and space above, the rest of the spots are orange-ochreous, the one at subcosta slightly 
whitish; the submarginal ocelli are faint; the marginal ochreous spots faint except those at 
hind angle. Hing wing, ground colour as fore wing, the basal black lines are thin, edged in 
white, those on the proximal side of the irregular white bar stronger, the white bar commencing 
at the subcosta fades out toward the inner fold which it faintly crosses above the anal angle; 
the postdiscal series of ochreous lunules, outlined black proximally, become olive above the anal 
angle; the admarginal pale lunules with black internally are faint, except toward the anal angle, 
where the black spots on an olive ground are stronger; marginal border orange, becoming olive 
toward the anal angle; edge greyish olive with thin whitish fringe. 



26o V. G. L. van SOMEREN 

Neallotype female. Malawi: Soche Mt., 5000 feet, 26.iv.1966 (A. J. Martin). 
To be deposited in the B.M.(N.H.). 

Variation (a). The specimen from the Mlosa Stream, besides being smaller, has on the upper- 
side, the hind-marginal marks of the fore wing bar strongly lilac ; the postdiscal orange spots 2-4 
less clear and the marginal ochreous spots limited to the two at the hind angle. On the hind 
wing, the discal patch is narrower and without any orange scaling in the postdiscal line; the 
golden ochreous lunules on the border narrower and less clear. On the underside there is a 
corresponding reduction in the pattern. 

Variation (b). Upperside. The female from the Nchisi Forest (26.iii.1967) is semi-erythristic 
in that the marks of the fore wing discal bar are strongly tinted with orange except for the three 
subcostal ones, but the mark in ia is lilac; the postdiscal spots, except for the two subapical 
are orange-rufous. The marginal spots are small with the exception of the two at the hind 
angle. On the hind wing the discal patch is reduced in width and more purplish blue on the 
lower half but is shaded with orange on the upper outer border. The submarginal blue spots 
are small and the marginal orange is narrow and broken. Underside. Fore wing, strongly 
rufescent, the basal area of ib blackish; the margin rusty. Hind wing, the discal area is 
brownish, while the discal bar is hardly indicated, except for a thin area of brownish; postdiscal 
marks are rusty, thinly black proximally and extending to above the anal angle ; the admarginal 
dark spots are faint at the upper angle, but become strong in the region of the tails and double 
in the anal angle; accentuated with white proximad; the marginal border dull orange shading 
to olive at the anal angle; edge olive-grey with narrow white fringe. 

Variation (c). Upperside. The second specimen from Nchisi (2.1.1967) is very like the 
neallotype on the fore wing, but the marginal orange spots are only visible at the hind angle 
where they coalesce. The hind wing discal patch is very narrow, slightly less shaded with bluish 
lilac on the borders; the submarginal lilac -blue spots distinct and the orange border strong. 
The underside, very similar to the type-specimen. 

There is some resemblance between this race and brevicaudatus of the southern 
Highlands of Tanzania, which has a wider, more uniform white bar in the female, 
and specimens of xiphares from the Nyika Plateau (Malawi-Zambia) appear to 
belong to brevicaudatus. 



Charaxes xiphares kilimensis ssp. n. 

(PI. 12, figS 96, 97) 

This new subspecies of Ch. xiphares belongs to the north-eastern group of the 
species which, at the present, includes brevicaudatus Schultze, maudei Joicey & 
Talbot, desmondi van Someren and kulal van Someren, in which the females exhibit 
a departure from che usual female pattern of the more southern group, most ot 
which have a large ochre patch in the disc of the hind wing. In the northern group 
the hind wing discal patch is white with strong blue scaling on the borders or overall. 
The fore wing discal bar is white, while the postdiscal spots, which are pronounced, 
are ochreous to orange-ochre. 

This new subspecies from western slopes of Mt Kilimanjaro exhibits characters 
which place it intermediately between brevicaudatus of the northern end of Lake 
Nyasa and the southern highlands of Tanzania, which has very short tails in both 
sexes, and maudei of the Usambara Mts and possibly the Ulugurus, which has very 
long tails in both male and female. 



REVISIONAL NOTES ON AFRICAN CHARAXES 261 

Male. Fore wing length 49 mm. Upperside. Fore wing, ground colour deep blue-black 
with strong blue sheen in side light, base of costa browner. Discal blue spots rather small, two 
spots just beyond end of cell, upper one a streak, spot below larger, spot sub-basal in 4 larger 
and round, spot below it in 3 more elongate but smaller with a minute dot beyond, no spot in 2 
and spot in ib small and round but mark in ia an elongate streak widest proximally and tapering 
toward postdiscal line. Postdiscal series: two subapical in 8-7 white, spots in 6-4 slightly 
smaller and blue, spot in 3 slightly larger, that in ib double. Margin without orange spots 
except for a slight indication in ib. Hind wing with a large discal bluish white patch not 
reaching beyond 5, with strong blue suffusion on the borders, represented at the subcosta by a 
discrete blue spot; in the postdiscal row there are two discrete blue spots in the upper sector, 
but there are confluent blue marks on the outer border of the discal patch, with black scaling in 
between. Submarginal blue spots distinct from 2-6; border of wing with orange-rufous lunules 
from anal angle to 6, edged with black. Tails black, of about equal length, 4 mm. Underside. 
Fore wing, ground colour earthy grey with a slight brassy tone, the whole with a satiny sheen 
in side light except in mid areas of 1 and 2, which are dull; the base of the wing olive crossed 
by narrow black lines outlined in white ; the discal spots represented in olive, proximally edged 
with black with a suggestion of white in between the postdiscal spots; the two upper subapical 
spots ochreous, the rest golden olive, the tornal mark olive to greyish distally with conspicuous 
double half-moon, black in centre, spot in 2 above with slight black distally. Margin with very 
obscure olive marks, more obvious in ib and 2. Hind wing ground colour as fore, the sub-base 
crossed with zigzag olive line narrowly edged in black; the distal zone more bronzy, with a zigzag 
narrow whitish line from costa to 2, a narrow black line through end of cell area. Postdiscal 
series of spots from costa to anal angle golden olive narrowly edged with black proximally, the 
mark in the anal angle a long crescent double edged in black; submarginal spots bluish grey, 
those toward the anal with black dot distally; marginal lunules golden olive, more greenish at 
anal angle; edge black with very narrow white fringe. Tails mostly black with olive mid line 
at base. 

Female. Fore wing length 55 mm. Upperside. Fore wing, ground colour purplish brown- 
black in basal triangle, blacker on distal half of wing. Disc of wing crossed by a broad white 
curved band extending from the costa to hind margin, consisting of 4 elongate white marks, 
including white area on costa, beyond end of cell followed by an almost oval spot sub-basad in 3, 
the mark in 2 more elongate somewhat pear-shaped, the white mark in ib double, small mark 
above that a blunted triangle, followed by a long streak in ia, the marks in ia, ib shaded 
with violet scales. Postdiscal spots distinct, a large subcostal subapical mark is whitish and 
rounded, spot in 6 smaller, those in 5-2 smaller and orange in colour, mark in ib double and 
contiguous with the discal marks. Margin with slight internervular orange marks, double in ib. 
Hind wing ground colour black in basal triangle, more purplish black in dark border; disc of 
wing with large violet-bluish patch, more whitish toward bases of 5-6, with a large whitish 
quadrate mark at subcosta in 8 ; the discal patch itself shaded with violet and on its distal border 
is a series of dyslegnic rounded orangish marks from 2-5. Submargin with a series of lilac-blue 
spots, distinct from anal angle to 5 ; margin with strong orange border of confluent lunules which 
shade to olive green at anal angle; edge black with narrow white fringe. Tails black, 5 mm long. 
Underside. Ground colour as in the male. Fore wing, basal triangle with series of strong black 
lines margined with bluish white, three crossing the cell and two at end of cell, with a black 
spot at sub-basal of ib and with a short bar in sub-base of 2. The discal white bar conspicuously 
outlined proximally in black; post-discal spot white in sub-apical area then increasingly orange 
to ib, with the double black spots in ib and 2 strongly marked; admargin with slight orange 
lunules strongest in ib. Hind wing ground colour as fore wing; fine black lines outlined in white 
at basal triangle; discal bar represented by a pale ochre-greyish band, dyslegnic on its outer 
border but edged internally by a narrow black and white zigzag line ; postdiscal series of rather 
indistinct ochreous lunules, shaded brownish in lower half, slightly more olive and more defined 
above anal angle; submarginal series of greyish lunules, broadly edged with black, distally 
touch the marginal orange-ochre lunules which shade to olive at the anal angle; extreme edge 
black. 



262 V. G. L. van SOMEREN 

Holotype male. Tanzania: Lower slopes of west Kilimanjaro at Maua Estate, 
September 1966. (Collector Edmund, for Major I. Grahame.) 

Allotype female. Taken in the same locality, February 1964 (A. F. Brown). 
Deposited in the B.M.(N.H.). 

Range: This subspecies is at present known only from the west Kilimanjaro area. 

Charaxes xiphares ludovici Rousseau-Decelle 
(PI. 12, fig. 95) 
Charaxes ludovici Rousseau-Decelle, 1933 : 2 7 I - 

This name was associated with subspecies brevicaudatus Schultze originally (van 
Someren, 1964 : 195) but was not formally synonymized. I am unable to form an 
opinion as to the validity of this race, but Major I. Grahame is of the opinion that 
it is sound. 

The unique type is from Lake Nyassa, E. Africa. (Grahame coll.) 

Charaxes xiphares desmondi van Someren 

Charaxes xiphares desmondi van Someren, 1964 : 201, pi. 8, fig. 54. 

It may be pointed out that the female specimen figured in the above paper is 
the neallotype described in the text. 

SYSTEMATIC LIST 

Charaxes xiphares (Cramer, 1781) 

Charaxes xiphares woodi van Someren, 1964. Type locality: Malawi. 

Range: Malawi. 
Charaxes xiphares kilimensis ssp. n. Type locality: Tanzania: West Kilimanjaro, 

Maua Estate. 
Charaxes xiphares ludovici Rousseau-Decelle, 1933. Type locality: Lake Nyassa. 
Charaxes xiphares desmondi van Someren, 1939. Type locality: S.E. Kenya, Teita 

Range. 

ACKNOWLEDGEMENTS 

I am greatly indebted to the many who have assisted me with specimens, photo- 
graphs and information. I wish especially to place on record my thanks to Mr T. G. 
Howarth, Mr D. E. Kimmins and Mr C. F. Huggins of the Department of Entomology, 
British Museum (Natural History), London, for constant help, reading through the 
typescript and mounting the plates. For the loan of material I am indebted to the 
late Dr R. M. Fox, of the Carnegie Museum, Pittsburgh, U.S.A.; Dr F. Rindge 
of the American Museum of Natural History, New York, U.S.A. ; Dr P. Viette of the 
Museum National d'Histoire Naturelle, Paris, France ; Dr H. J. Hannemann of the 
Humboldt University Museum, Berlin, E. Germany; Monsieur J. Plantrou of Paris, 



REVISIONAL NOTES ON AFRICAN CHARAXES 263 

France; Major I. Grahame of Lamarsh, Suffolk, England; Dr C. H. McCleery of 
Zomba, Malawi; Mr P. T. Martin of Limbe, Malawi; Mr E. Taylor of the Hope 
Dept. of Entomology, University Museum, Oxford, England; Dr R. H. Carcasson, 
late of the National Museum, Nairobi, Kenya; the late Mr T. H. E. Jackson of 
Kitale, Kenya; and Dr A. H. B. Rydon, of North Chailey, Sussex, England, lor 
making genitalia preparations. 



REFERENCES 

References not given here will be found in Parts I-VTI of this revision. 

Aurivillius, C. 1889. En ny art slagtet Charaxes Ochs. Ent. Tidskr. 10 : 191-192. 
Butler, A. G. 1874. Descriptions of three new species and a new genus of Diurnal Lepidop- 

tera, from the collection of Andrew Swanzy, Esq. Trans. R. ent. Soc. Lond. 7 : 531-533, 

pi. 11. 

1887. On two small Collections of African Lepidoptera recently received from Mr H. H. 

Johnston. Proc. zool. Soc. Lond. 1887 : 567-574. 

Capronnier, J. B. 1889. Liste des Lepidopteres captures au Congo par Messieurs Thys, 

Legat, Martini et Macahdo en 1887. Annls Soc. ent. Belg. 33 : cxviii-cxxvi. 
Crowley, P. 1890. On some new species of African Diurnal Lepidoptera. Trans. R. ent. 

Soc. Lond. 1890 : 551-556, pis 17-18. 
Distant, W. L. 1879. On some African Species of Lepidoptera belonging to the Subfamily 

Nymphalinae. Proc. zool. Soc. Lond. 1879 : 703-709, pi. 54. 
Doubleday, E. 1844. List of the Specimens of Lepidopterons Insects in the Collection of the 

British Museum. 1 : 1-150. London. 
Doumet, N. 1861. Description de Lepidopteres nouveaux. Revue Mag. Zool. (2) 13 : 171— 

178, pi. 5, figs 1-2. 
Fabricius, J. C. 1793. Ent. Syst. 3 (1) : 1-487. Hafniae. 
Hewitson, W. C. 1859. Illustrations of New Species of Exotic Butterflies, 2 : 1-124, 60 pis., 

London. 

1876. Illustrations of New Species of Exotic Butterflies, 5 : 1-208, 60 pis. London. 

Holland, W. J. 1886. Contributions to a Knowledge of the Lepidoptera of West Africa. 

Trans. Am. ent. Soc. 13 : 325-332, pi. 8. 

1917. Two New West African Rhoplaocera. Ann. Carnegie Mus. 11 : 14-18, pi. 4. 

Howarth, T. G. 1969. Some African Nymphalidae (Lepidoptera). Proc. R. ent. Soc. Lond. 

(B) 38 : 141-156, pi. 11. 
Joicey, J. J. & Talbot, G. 1921. New forms of Rhopalocera. Bull. Hill Mus. Witley 

1 : 44-103, pis 5-16. 

1927. Four new Butterflies from the Island of St Principe. Entomologist 60 : 12-16. 

Karsch, F. 1892. Vorlaufige Beschreibung von drei neuen Lepidopteren von Bismarckburg 

im Togolande (Deutschwestafrika) . Ent. Nachr. 18 : 113-117. 
Le Cerf, F. 1927. Description d'un Charaxes nouveau d'Abyssinie. Encycl. ent. B.3, 

Lep. 2 : 144. 
1932. Charaxes nouveaux du Congo Beige (Lepid. Rhopal.). Bull. Mus. natn. Hist. nat. 

Paris (2) 4 : 405-406. 
Rothschild, W. 1894. Some new Species of Lepidoptera. Novit. zool. 1 : 535-540, pi. 12. 
1902. Some New N.E. African Lepidoptera discovered by Oscar Neumann. Novit. zool. 

9 : 595-598. 
Rousseau-Decelle, G. 1956. Note sur une sous-espece nouvelle de Charaxes africain. 

Bull. Soc. ent, Fr. 61 : 91-92, pi. 1. 
Stoneham, H. F. 1931. A New Form of Charaxes numenes, Hew., from Kenya Colony. 

Bull. Stoneham Mus. No. 7. 






264 



V. G. L. van SOMEREN 



van Someren, V. G. L. 1963. Revisional Notes on African Charaxes (Lepidoptera : Nym- 
phalidae). Part I. Bull. Brit. Mus. nat. Hist. (Ent.) 13 (7) : 195—242, 19 pis, 5 text-figs. 

1964. Part II. Bull. Brit. Mus. nat. Hist. (Ent.) 15 (7) : 181-235, 2 3 P' s . 4 maps. 

Bull. Brit. Mus. nat. Hist. (Ent.) 18 (3) : 
Bull. Brit. Mus. nat. Hist. (Ent.) 18 (9) 
Bull. Brit. Mus. nat. Hist. (Ent.) 23 (4) 



- 1966. Part III. 

- 1966a. Part IV. 

- 1969. Part V 
8 maps. 

- 1970. Part VI. 

- 1971. Part VII. 



45-100, 16 pis, 5 maps. 

: 277-316, 9 pis, 4 maps. 

: 75-166, 29 pis, 31 text-figs, 



Bull. Brit. Mus. nat. Hist. (Ent.) 25 (5) : 
Bull. Brit. Mus. nat. Hist. (Ent.) 26 (4) 



Ward, C. 1871. Description of New Species of African Diurnal Lepidoptera. 

mon. Mag. 8 : 1 18-122. 
Watkins, H. T. G. 1923. Notes on the Butterflies of the Banks Collection. 

56 : 204-209. 



197-249, 11 pis, 6 maps. 
181-225, IX pl s » 6 maps. 

Entomologist' s 



Entomologist 



INDEX 

Synonyms in italics 



aequatorialis, 244 
albimaculatus, 252 
albipunctatus, 225 
amelia, 228 
amelina, 231 
atribasis, 255 
aubergeri, 256 

barnesi, 253 
bipunctatus, 246 
brunneus, 235 
bubastis, 251 
busogus, 234 

caeruleipunctatus, 227 

desmondi, 262 
dux, 231 

fuscus, 240 

gabonica, 221 
galba, 218 

hadrianus, 221 
hildebrandti, 218, 219 

imperialis, 223 

jacksonianus, 234 
johnsoni, 246 

katangensis, 219 
kilimensis, 260 



lactetinctus, 232 
laticatena, 243 
lecerfi, 232 
ludovici, 227, 262 
manica, 255 
marica, 236 
marginatus, 239 
mccleeryi, 257 
mixtus, 250 
neumanni, 245 
numenes, 241 
obsolescens, 244 
paulianus, 227 
pseudosmaragdalis, 256 
regius, 228 
subrubidus, 255 
talagugae, 218 
thysii, 219 
tiridates, 235, 236 
tristis, 237 
ugandensis, 248 
ugandicus, 225 
ungemachi, 235 
victoriae, 231 
xiphares, 259 
woodi, 259 



Dr V. G. L. van Someren, 
The Sanctuary, Ngong, 
P.O. Box 24947, 
Karen, Kenya. 



PLATE i 

Charaxes 

Upper and undersides 
Fig. i. hildebrandti hiblderandti Dewitz, $. 
Fig. 2. hildebrandti hildebrandti Dewitz, $. 
Fig. 3. hildebrandti katangensis Talbot, o* (Congo: Katanga, Lulua) (F. G. Overlaet). Photos 

I. Grahame. 
Fig. 4. thysii Capronnier, § (French Equatorial Africa) (Jackson coll.). 

Fig. 5. hadrianus hadrianus Ward, $ (Congo: Katanga, Kafakumba). Photos I. Grahame. 
Fig. 6. hadrianus hadrianus Ward, $ (Congo: Katanga, Kafakumba). Photos I. Grahame. 
Fig. 7. hadrianus lecer\ Lathy, $ (Ghana Forest) (L. R. Cole). Photos B.M.(M.H.) Nos 

51274-5. 
Fig. 8. hadrianus lecer\ Lathy, $ (Nigeria: Warri) (Rothschild coll.). Photos B.M.(N.H.) 

Nos 50120-1. 
Fig. 9. hadrianus ? ssp. n. $ (Sierra Leone: Kanbui Hills, Bamba-wo) (G. D. Field). Photos 

B.M.(N.H.) Nos 512-667. 



Bull. Br. Mus. nat. Hist. (Ent.) 27, 4 



PLATE 1 




PLATE 2 

Charaxes 

Upper and undersides 

Figs 10 & ii. imperialis Butler, $ Type (Gold Coast [Ghana]). Photos B.M.(N.H.) Nos 

50132-3. 
Figs 12 & 13. imperialis Butler, $ Type (Gold Coast [Ghana]). Photos B.M.(N.H.) Nos 

50138-9. 
Fig. 14. imperialis Butler, <$ (Nigeria: Ikom). 

Fig. 15. imperialis Butler, $ (Ivory Coast: Banco- Abidjan). Photos J. Plantrou. 

Figs 16 & 17. albipunctus Joicey & Talbot, £ Paratype (Cameroun). Photos B.M.(N.H.) 

Nos 50134-5. 



Bull. Br. Mus. nat. Hist. (Ent.) 27, 4 



PLATE 2 




PLATE 3 

Charaxes 

Upper and undersides 

Figs 18 & 19. albipunctus Joicey & Talbot, ? Allotype (Cameroun). Photos B.M.(N.H.) 

Nos 50136-7. 
Fig. 20. imperialis, S transitional to albipunctus Joicey & Talbot (Uganda: Bwamba 

Valley, E. side Semliki River). 
Fig. 21. imperialis, $ transitional to albipunctus Joicey & Talbot (Uganda: Bwamba 

Valley, E. side Semliki River). 
Fig. 22. imperialis, $ form caerulipunctus forma n. Holotype (Uganda: Bwamba Valley) 

(I. Grahame coll.). Postdiscal spots fore wing blue; hind wing blue patch 

extended towards inner fold, costal spots large and bluish white. . 
Fig. 23. ugandicus ssp. n., <$ Holotype (Uganda: Masaka district, Katera Forest, W. 

shore Lake Victoria) (van Someren coll.). 
Fig. 24. ugandicus ssp. n., $ Allotype (Uganda: Masaka district, Katera Forest, W. shore 

Lake Victoria) {van Someren). 



Bull. Br. Mus. nat. Hist. (Ent.) 27, 4 



PLATE 3 




PLATE 4 

Charaxes 

Upper and undersides 

Fig. 25. imperialis paulianus Rousseau-Decelle, $ Type (Congo: Katanga, Kafakumba). 

Photos J. Plantrou. 
Fig. 26. imperialis paulianus Rousseau-Decelle, $ Type (Congo: Katanga, Kafakumba). 

Photos J. Plantrou. 
Fig. 27. imperialis ludovici Rousseau-Decelle, c? (Zambia: Mwinilunga) (C. B. Cottrell). 

Fig. 28. imperialis ludovici Rousseau-Decelle, $ (Zambia: Mwinilunga) (C. B. Cottrell). 

Fig. 29. ameliae ameliae Doumet, £ (Ivory Coast). 

Fig. 30. ameliae ameliae Doumet, ? (Ivory Coast). 

Figs 31 & 32. ameliae victoriae ssp. n., <J Holotype (Uganda: Masaka district, Katera Forest. 

W. shore Lake Victoria) {van Someren). 



Bull. Br. Mus. nat. Hist. (Ent.) 27, 4 



PLATE 4 





PLATE 5 

Charaxes 

Upper and undersides 

Figs 33 & 34. ameliae victoriae ssp. n., $ Allotype (Uganda: Masaka district, Katera Forest, 

W. shore Lake Victoria) {van Someren). 
Fig. 35. ameliae victoriae ssp. n., <$ Paratype (Uganda: Masaka district, Katera Forest, 

W. shore Lake Victoria) {van Someren). 
Fig. 36. ameliae victoriae ssp. n., <j> Paratype (Uganda: Masaka district, Katera Forest, 

W. shore Lake Victoria) {van Someren). 
Fig. 37. ameliae amelina Joicey & Talbot, <? (Malawi: Nkata Bay, Lake Nyasa). 

Fig. 38. ameliae amelina Joicey & Talbot, ? (Malawi). 

Fig. 39. ameliae amelina Joicey & Talbot, ? (Malawi). 

Fig. 40. lactetinctus lactelinctus Karsch, £ Holotype (Togo). Photos B.M.(N.H.) Nos 

50140-1. 



Bull. Br. Mus. nat. Hist. (Ent.) 27, 4 




PLATE 5 




PLATE 6 

Char axes 

Upper and undersides 

Fig. 41. lactetinctus Karsch, $ (Gold Coast [Ghana]). 

Fig. 42. lactetinctus Karsch, ? (Nigeria). 

Figs 43 & 44. busogus ssp. n., cJ Holotype (Kenya: Busoga district, Broderick Falls Scarp) 

(van Someren). 
Figs 45 & 46. busogus ssp. n., $ Allotype (Uganda: Metu Hills, N. W. Madi, W. Nile, v-vi.1954) 

(van Someren) . 
Figs 47 & 48. busogus $ form jacksonianus van Someren (N.W. Kenya). 



Bull. Br. Mus. nat. Hist. (Ent.) 27, 4 



PLATE 6 






PLATE 7 

Charaxes 

Upper and undersides 

Fig. 49. lactetinctus ungemachi, <J form brunneus Carpenter (Ethiopia: Youbdo). 

Fig. 50. lactetinctus ungemachi Le Cerf, $ Paratype (Ethiopia: Youbdo). 

Fig. 51. tiridates tiridatinus Rober, $ (Uganda). 

Fig. 52. tiridates tiridates Cramer, <J (Ivory Coast). 

Fig. 53. tiridates tiridatinus Rober, $ (Uganda). 

Fig. 54. tiridates tiridates Cramer, $ (Ivory Coast). 

Fig. 55. tiridates tiridatinus Rober, S (S.W. Uganda: Kayonza). 

Fig. 56. tiridates tiridatinus Rober, $ (N.W. Kenya). 



Bull. Br. Mus. nat. Hist. (Ent.) 27, 4 



PLATE 7 




Fig. 


57- 


Fig. 


58. 


Fig. 


59- 


Fig. 


60. 


Figs 6i & 62. 


Fig. 


63- 


Fig. 


64. 



PLATE 8 

Charaxes 

Upper and undersides 

tiridates marginatus Rothschild, $ Holotype (Ethiopia: Sheko, 25.iv.1901). 

tiridates marginatus Rothschild, <$ (Ethiopia). 

fuscus Plantrou, <J Holotype (Central African Republic: Bangui, ix.1966) 

(R. P. Godart). 

numenes numenes Hewitson, £ (Ivory Coast). 

numenes numenes Hewitson, 2 $ (Ivory Coast). 

numenes Hewitson, $ intermediate cline (Central African Republic: Bangui). 

numenes Hewitson, $ intermediate cline (Central African Republic: Bangui). 



Bull. Br. Mus. nat. Hist. (Ent.) 27, 4 



PLATE 8 




Fig. 


65- 


Fig. 


66. 


Fig. 


67. 


Fig. 


68. 


Fig. 


69. 


Fig. 


70. 


Fig. 


7i- 


Fig. 


72. 



PLATE 9 

Charaxes 

Upper and undersides 

numenes aequatorialis ssp. n., $ Holotype Uganda: Kayonza, Kigezi, 6.ix.i952) 

(van Someren). 

numenes aequatorialis ssp. n., $ Allotype (Uganda: Kayonza Forest, Kigezi, v-vi. 1957) 

(van Someren). 

numenes neumanni Rothschild, o* Holotype (Wori-Gamitscha Kafla, 5.xi.i90i). 

Photos B.M.(N.H.) Nos 49045-6. 

numenes neumanni Rothschild, <£ (Ethiopia: Lalo Kel) (Joicey Bequest). Photos 

B.M.(N.H.) Nos 46086-7. 

bipunctatus bipunctatus Rothschild, <£ Holotype (Ghana). Photos B.M.(N.H.) 

Nos 49040-1. 

bipunctatus bipunctatus Rothschild, $ Allotype (Ghana). Photos B.M.(N.H.) 

Nos 49038-9. 

bipunctatus bipunctatus Rothschild o* (Ghana). 

bipunctatus Rothschild, $ (Ghana). 



Bull. Br. Mus. nat. Hist. (Ent.) 27, 4 



PLATE 9 




PLATE 10 

Charaxes 

Upper and undersides 

Fig. 73. bipunctatus ugandensis ssp. n., g Holotype (Uganda: Toro Forest (J.) x.1949) 

(van Someren). 
Fig. 74. bipunctatus ugandensis ssp. n., $ Allotype (Uganda: Toro, Kibali Forest, v-vi.1956) 

(van Someren). 
Fig. 75. bipunctatus ugandensis ssp. n., $ Paratype (S.W. Uganda: Kayonza Forest). 
Fig. 76. mixtus Rothschild, £ Holotype (Lokolele, mid Congo River). Photos B.M.(N.H.) 

Nos 46082-3. 
Fig. 77. mixtus Rothschild cj. 
Fig. 78. mixtus Rothschild (J. 
Fig. 79. mixtus Rothschild $. 
Fig. 80. bubastis Schultze <J (Cameroun: Bipindi) B.M.(N.H.). 



Bull. Br. Mus. nat. Hist. (Ent.) 27, 4 



PLATE 10 




PLATE ii 

Charaxes 

Upper and undersides 

Fig. 8i. albimaculatus ssp. n., <J Holotype (Congo: Stanleyville) (Ertl. coll. ex Joicey 

Bequest) B.M.(N.H.). 
Fig. 82. albimaculatus ssp. n., 9 Allotype (S. Congo: Kafakumba, Katanga) (Overlaet 

coll. ex Le Moult, in coll. J. Plantrou). 

barnesi Joicey & Talbot, S (Island of Principe, Gulf of Guinea, iv-v.1926). 

Photos B.M.(N.H.) Nos 46078-9. 

barnesi Joicey & Talbot, $ (Island of Principe, Gulf of Guinea iv-v.1926). 

Photos B.M.(N.H.) Nos 46084-5. 

manica subrubidus ssp. n., cJ Holotype (Congo: Brazzaville, x.1966) (J. Plantrou). 

manica subrubidus, $ form atribasis forma n., Holotype (Congo: Brazzaville, 

19.ix.1966) (J . Plantrou) . 



Fig. 


83. 




Fig. 


84. 




Figs 85 
Figs 87 


& 86. 
& 88. 



Bull. By. Mus. nat. Hist. (Ent.) 27, 4 





83 v ^ 






Figs 89 & 


90. 


Figs 


. 91 & 


92. 


Fig. 


93- 




Fig. 
Fig. 


94- 
95- 




Fig. 
Fig. 


96. 
97- 





PLATE 12 

Charaxes 

Upper and undersides 

manica subrubidus, ? form aubergeri forma n., Holotype (Congo: Brazzaville, 

25. v. 1967) (/. Plantrou). 

manica subrubidus, $ form pseudosmaragdalis van Someren & Jackson. 

mccleeryi sp. n., $ Holotype (Tanzania: Bundaki Hills, Uluguru Mts, 6000 ft, 

vii.1966) (C. H. McCleery). 

mccleeryi sp. n., ? Allotype (Tanzania: Bunduki Hills, Uluguru Mts, 6000 ft, 

vii.1966) (C. H. McCleery). 

xiphares ludovici Rousseau-Decelle, $ Type (Malawi: Lake Nyasa) (I. Grahame 

coll.). 

xiphares kilimensis ssp. n., <J Holotype (Tanzania: Lower slopes W. Kilimanjaro, 

Maua Estate, ix.1966) {Edmund). 

xiphares kilimensis ssp. n., ? Allotype (Tanzania: Lower slopes W. Kilimanjaro, 

Maua Estate, ii.1964) [A. F. Brown). 



3ull. Br. Mus. nat. Hist. (Ent.) 27, 4 



PLATE 12 






A LIST OF SUPPLEMENTS 

TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 
18 plates, 270 text-figures. August, 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 : 500 text-figures. September, 

1965- £3-25. 

5. Ahmad, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156: 
475 text-figures. November, 1965. (out of print) £2.15. 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. 

£3-15- 

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 
text-figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera) . Pp. 509. £8.50. Reprinted 1972. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322: 348 text-figures. August, 1967. £8. 

11. Mound, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172: 
82 text-figures. May, 1968. £4. 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. 
November, 1968. £5. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North- Western Europe 
(Hymenoptera : Chalcidoidea) . Pp. 908: 686 text-figures. November, 1969. 

£*>■ 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 

68 plates, 15 text-figures. October, 1971. £12. 

18. Sands, W. A. The Soldierless Termites of Africa (Isoptera Termitidae). 
Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. 



33EIS Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 



ON EUROPEAN PTEROMALIDAE 

(HYMENOPTERA) : 

A REVISION OF CLEONYMUS, 

EUNOTUS AND SPANIOPUS, 

WITH DESCRIPTIONS OF NEW 

GENERA AND SPECIES 



Z. BOUCEK 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 27 No. 5 

LONDON: 1972 



ON EUROPEAN PTEROMALIDAE [ 3 o octwz 

(HYMENOPTERA): 

\ REVISION OF CLEONYMUS, EUNOTUS AND 

SPANIOPUS, WITH DESCRIPTIONS OF NEW 

GENERA AND SPECIES 



BY 

ZDENEK BOUCEK 

Commonwealth Institute of Entomology 



Pp 265-315; 62 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 27 No. 5 

LONDON: 1972 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 27, No. 5 of the Entomological 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Ent.). 



Trustees of the British Museum (Natural History), 1972 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 18 October, 1972 Price £1.85 



ON EUROPEAN PTEROMALIDAE 

(HYMENOPTERA): 

A REVISION OF CLEONYMUS, EUNOTUS AND 

SPANIOPUS, WITH DESCRIPTIONS OF NEW 

GENERA AND SPECIES 

By Z. BOUCEK 



CONTENTS 



Synopsis 
Introduction 
Acknowledgements 
Cleonyminae . 

Cleonymus Latreille 
Eunotinae 

Eunotus Walker 
Miscogasterinae 

Susteraia gen. n. 

Semiotellus Westwood 
Pteromalinae 

Veltrusia gen. n. 

Strejcekia gen. n. 

Rhizomalus gen. n. 

Pteromalus Swederus 

Peridesmia Forster 

Spaniopus Walker 
References . 
Parasite index 



Page 
267 
267 
268 
269 
269 
274 
274 
287 
287 
290 
292 
292 

295 
298 
302 
303 
305 
314 
315 



SYNOPSIS 



This paper supplements Graham's recent work (1969) in revising the European species of 
several genera of Pteromalidae and giving descriptions of new taxa. Most of the 14 new 
species are from central and southern Europe. In Cleonymus three species are recognized 
including two new ones, in Eunotus nine species (two new), in Spaniopus seven species (two 
new). Also described are five new species belonging to four new genera, one of them in the 
subfamily Miscogasterinae, the other three in the Pteromalinae. In Peridesmia one new 
species is added to the two previously known; similarly one species is described in Semiotellus 
and one in Pteromalus. The study of extensive fresh material and of the relevant types results 
in better recognition of some species and in the establishment of four new specific synonyms. 

INTRODUCTION 

The magnificent and most useful monograph of the Pteromalidae of north-western 
Europe by Graham (1969) covers the Pteromalid fauna of Europe to a greater 
extent than its title would suggest. Where possible, it provides keys to the Euro- 
pean genera and species. 



268 Z. BOUCEK 

In some groups, apparently, most species have already been described but in some 
other groups information is still very incomplete. It is mainly in the latter groups 
that rich material has been collected together and the present contribution is the 
result of its study. This paper may therefore be regarded as a supplement to 
Graham's work. 

The taxa are classified in the same way and described mainly in the same form. 
The morphological terms are also used mainly in the same sense as Graham, except 
for the following. 

Prepectus is used instead of postspiracular sclerite, for in the more primitive 
Chalcidoids it forms an unbroken belt in front of the mesopleurae, so that 'post- 
spiracular sclerite', pointing to an independent sclerite on each side of the thorax 
(as it occurs generally in more apomorph forms) does not seem quite as appropriate 
as the older term 'prepectus'. 

The mesopleura of the thorax is divided by the oblique pleural line running from 
the base of the mid coxa towards the base of the fore wing (where it branches) into 
the anteroventral mesepisternum and posterodorsal mesepimeron. The mesepimeron 
is usually subdivided into the upper and lower epimeron. Graham calls only the 
lower part epimeron and considers the upper, frequently smooth part, as belonging 
to the mesepisternum, following Thomson. 

The gastral tergites are counted, as Graham does in the text, i.e., the basal, post- 
petiolar tergite as the first, the spiracle-bearing tergite as the sixth. 

The following abbreviations are used for depositories of the collections : 

BMNH — British Museum (Natural History), London. 

IEE, Madrid — Instituto Espanol de Entomologia, Madrid, Spain. 

MCSN, Genoa — Museo Civico di Storia Naturale, Genoa, Italy. 

MHN, Geneva — Museum d'Histoire Naturelle, Geneva, Switzerland. 

MNHN, Paris — Museum National d'Histoire Naturelle, Paris, France. 

MNHU, Berlin — Museum fiir Naturkunde der Humboldt-Universitat, Berlin, 
East Germany. 

NM, Dublin — National Museum of Ireland, Dublin. 

NM, Prague — Narodni Museum, Prague, Czechoslovakia. 

NM, Vienna — Naturhistorisches Museum, Vienna, Austria. 

TM, Budapest — Termeszettudomanyi Muzeum, Budapest, Hungary. 

USNM, Washington— U.S. National Museum, Washington, U.S.A. 

UZI, Lund — Universitetets Zoologiska Institution, Lund, Sweden. 

UZM, Copenhagen — Universitetets Zoologiske Museum, Copenhagen, Denmark. 

ZI, Leningrad — Zoological Institute of the Academy of Sciences of the U.S.S.R., 
Leningrad. 

ZIPF, Zagreb — Zooloski Institut Poljoprivrednog Fakulteta, Zagreb, Yugo- 
slavia. 

ACKNOWLEDGEMENTS 

I wish to express my thanks to the Keeper and staff of the Department of Ento- 
mology of the BMNH for the opportunity of research and access to collections. My 
thanks are also extended to various colleagues from different institutions, mainly 



EUROPEAN PTEROMALIDAE 269 

for assistance in submitting the types or other material for study, in particular the 
following: Mrs E. Mingo (IEE, Madrid), Mr 0. Bakkendorf (UZM, Copenhagen), 
Dr C. Besuchet and Dr I. Lobl (MHN, Geneva), Dr M. Fischer (NM, Vienna), 
Dr M. W. R. de V. Graham (Oxford), Dr K.-J. Hedqvist (Stockholm), Prof. C. 
Lindroth (and Staff, UZI, Lund), Dr J. Papp and Prof. G. Szelenyi (TM, Budapest), 
Dr V. A. Trjapitzin (ZI, Leningrad), Dr D. J. Williams (Commonw. Inst, of Ent., 
London, for indentifications of Coccids) and Mr W. G. Tremewan (BMNH), for 
linguistic help. 



CLEONYMINAE 
CLEONYMUS Latreille 

Cleonymns Latreille, 1809 : 29. Type-species: Diplolepis depressa (Fabricius) ; designated by 
Latreille, 1810. 

The subsequent references are quoted by Graham (1969) and are not repeated 
here. 

Kerrich & Graham (1957) and Graham (1969) recognize two European species, 
C. laticornis Walker and C. obscurns Walker. A study of further material revealed 
two new species in southern Europe and threw doubts on the validity of C. obscurus. 

Key to European Species 

1 Frons in front of ocelli deeply regularly punctured, with narrow smooth interspaces 

between punctures (Text-fig. 4) ; vertex 1 -36-1 -48 times as broad as the relatively 
small eye. Fore wing in $ with hairs uniformly dark and long, even on the sub- 
hyaline spot below marginal vein (Text-fig. 2) ; infuscation of wing strong below 
postmarginal and stigmal vein but weak or absent below parastigma ; marginal vein 
fully 1 -g times as long as the stigmal. Fifth tergite in 9 in middle more than twice 
as long as the fourth and itself about 0-9 times as long as broad, coarsely raised- 
reticulate except along hind margin. Lateral ocellus about 1 -6 times its diameter 
from eye margin. <J unknown. (Balkan Peninsula) . balcanicus sp. n. (p. 270) 

- Frons sculpture different, either shagreened or rugulose; eyes usually larger, often as 
broad as the vertex. Fore wing in $ at least with some dense white hairs on 
hyaline spot below marginal vein, the latter at most 1 -72 times as long as the 
stigmal. Fifth tergite in the middle at most i-6 times as long as the fourth and 
itself at most 08 times as long as broad, its sculpture rather weak. Ocelli relatively 
larger, the lateral one at most about 1-15 its diameter from eye .... 2 

2 9. Body stout, gaster at most about twice as long as broad (Text-fig. 5), distinctly 

broader than the thorax, only 0-9-1 -16 times as long as head plus thorax combined. 
Head in dorsal view 2-2-15 times as broad as long, frons above shiny, shagreened 
(Text-fig. 9), piliferous punctures small. Scutellum usually transverse. Marginal 
vein less than i-6 times as long as the stigmal. Fifth tergite at least twice as 
broad as long in the middle. <$. Marginal vein hardly longer than the post- 
marginal, at most 1 -5 times as long as the stigmal . . . brevis sp. n. (p. 272) 



270 z. bouCek 

- $. Body slender, gaster at least 24 times as long as broad, at most only slightly 
broader than (1-02-1.1, if flattened then at most 1-2 times as broad as) the thorax. 
Head in dorsal view at least 2-2 times as broad as long, frons in front of ocelli very 
densely irregularly rugulose, usually dull. Scutellum usually slightly elongate. 
Marginal vein at least 1 7 times as long as the stigmal. Fifth tergite less trans- 
verse, o*. Marginal vein distinctly longer than the postmarginal and 1 -8-i -9 
times as long as the stigmal . laticornis Walker (and obscurus Walker) (p. 283) 

Cleonymus balcanicus sp. n. 

(Text-figs 1-4) 

$. Head and thorax mainly cupreous, in places sometimes with a weak bluish or greenish 
tint; propodeum and gaster bluish black or bronzy black, hind coxae dark green dorsally, 
bright cupreous laterally. Antennae and legs mainly dark rufous, but pedicel, first flagellar 
segment, preclava and clava more or less inf uscate ; fore tibia sometimes externally with bluish 
tint, mid and hind tibiae dorsally more or less inf uscate ; tarsi pale testaceous, inf uscate apically. 
Fore wing with an extensive brown cloud on disc extending to postmarginal vein and basad 
usually as far as below parastigma (Text-fig. 2) ; a small infuscation indicated also along hind 
margin in second quarter of wing. Hind wing subhyaline. Length 4-4-5-6 mm (the latter 
the holotype) . 

Head distinctly broader than mesoscutum (as 1-2-1-25 to 1), dorsally about 2-3-2-4 times as 
broad as long, with inner eye-margins slightly diverging forward. POL : OOL about as 
1-7 to 1. Head in facial view (only head capsule measured, excluding mandibles) 1-42-1.44 
times as broad as high. Ocelli relatively small, the lateral one more than 1 -5 times its diameter 
from eye. Frons in front of ocelli simply reticulate-punctured, interspaces narrow but distinct, 
smooth (Text-fig. 4) ; face further down at sides with meshes more lengthened, slightly rugose- 
reticulate; the same in transverse sense on vertex behind ocelli; upper frons not distinctly 
angulately or sculpturally separated from the lower (more vertical) part, scrobes also not 
indicated. Relative measurements: head width 82, height 57, frons (minimum distance 
between eyes) 41, eye 36-5 : 30, malar space 23, mouth width 33, scapus length 24, flagellum 
plus pedicellus combined about 67. Antenna similar to that of C. laticornis, only slightly 
stouter, all segments between pedicel and clava moderately transverse (in laticornis fourth 
flagellar segment usually slightly longer than broad) ; pedicellus as long as two following seg- 
ments combined, the first of them shorter than the second; processus of preclava broad and 
only about as long as the body of preclava. 

Dorsum of thorax with relatively coarse, deep and fairly regular reticulation-punctation 
and rather coarse but uniform whitish pubescence (as in C. laticornis). Scutellum about i-i 
times as long as broad, feebly convex, reticulation on its disc shallower than that on mesoscutum, 
axillar furrows anteriorly only moderately deep. Propodeum in the middle 0-45-0-5 times as 
long as scutellum, the median part superficially very weakly reticulate, much as in laticornis, 
but posterior flange laterad of nucha in dorsal view not quite as broad as the crenulate furrow 
basally at metanotal margin. Metapleura not very densely hairy. Mesepimeron in middle 
with deep fovea as a part of the curved groove separating the reticulate lower epimeron, the 
fovea not reaching metanotal margin; upper mesepimeron nearly smooth on disc but finely 
rugulose at sides, in particular anteriorly. Fore wing about 2 -7 times as long as broad, infumate 
all over, with darker brown markings as given in the key (Text-fig. 2), its pubescence relatively 
rough, dark, not paler on pale parts of the wing blade. Relative measurements (on the holo- 
type) : costal cell 90 : 10, marginal vein 44, postmarginal vein 37, stigmal vein 23, maximum 
distance between upper edge of stigma and the postmarginal vein 6-6. Marginal vein 1-9-1-96 
times as long as the stigmal. 

Gaster 1-1-1-2 times as long as head plus thorax combined, about 2-6 times as long as broad 
itself, distinctly flattened dorsally, about i-i times as broad as thorax, broadest just behind 
middle, i.e., at basal half of fifth (postpetiolar) tergite. First tergite mainly smooth, following 



EUROPEAN PTEROMALIDAE 



271 



tergites with a smooth belt along hind margin. Otherwise gaster dorsally reticulate, very 
coarsely so on the fifth tergite. This tergite is the largest (Text-fig. 1), nearly as long as tergites 
1 to 4 taken together (in median line), 0-89-0-95 times as long as broad itself. Hind margins 
of first, fourth and fifth tergites broadly shallowly emarginate. Hypopygium hardly reaching 
one-third of gaster. Hairs of gaster anteriorly whitish, posteriorly blackish, 
c?. Unknown. It might be recognizable by the sculpture of head. 

Biology not known. 

Holotype $. Greece: Attiki, Cape Sunion, SE of Athens, 7.1V. 1917 (Fodor); in 
TM, Budapest. 

Paratype. Bulgaria: i $, 7.V11.1964 (Strejcek); in Boucek Collection. 




Figs 1-9. Cleonymus. 1-4. C. balcanicus, 1, gaster of $; 2, part of fore wing with pilosity 
and infuscation indicated; 3, head in lateral view; 4, sculpture on frons in front of median 
ocellus. 5-9. C. brevis, 5, gaster of $; 6, fore wing of ?; 7, head in lateral view; 8, head 
of $ with antenna; 9, type of sculpture in front of median ocellus. 



272 z. bouCek 

Cleonymus brevis sp. n. 

(Text-figs 5-9) 

$. Body mainly dark (though in places bright) green to bronzy, more or less with bright 
violaceous reflections on vertex posteriorly, on sides of pronotum, on mesoscutum in front of 
axillae, on propodeum (at least laterally), on dorsal and lateral sides of hind coxae and anteriorly 
on first tergite. Vertical part of frons bright cupreous. Antennal flagellum dark brown to 
black, scape, and also femora, tibiae and tarsi of legs bright red, but tibiae mostly infuscate, as 
well as tarsi apically. Fore wing (Text-fig. 6) with a broad brown infumation between base 
of parastigma and apical fifth of wing blade, interrupted partly by large hyaline macula an- 
teriorly below marginal vein. Length 3-4-2 mm. 

Head 1 •1—1*14 times as broad as mesoscutum, in dorsal view 2*02-2*15 times as broad as 
long, with inner eye margins parallel; POL about 2*2-2*4 times the OOL. Head in facial 
view 1*37-1*38 times as broad as high. Frons between ocelli and the transverse blunt ridge 
(which is about 2*5 diameters from ocellus) fairly shiny, not very densely beset with piliferous 
punctures, interspaces generally broader than punctures but with an extremely fine broad- 
meshed engraved reticulation (alutaceous -f- puncturation = shagreening; Text-fig. 9). Sub- 
vertical part of frons angularly changing slope with the upper subhorizontal part (Text-fig. 7) , 
densely transversely striate-reticulate ; scrobes indistinct. Posterior part of vertex and occiput 
rugulose-reticulate. Relative measurements: head width 67, height 49, maximum distance 
between eyes 29, eye 34 : 29, malar space 19, mouth width about 28, scapus length 23, flagellum 
plus pedicellus combined 58. Pedicellus slightly longer than two following segments com- 
bined, these segments slightly transverse, subequal in length, the first narrower than the 
second; middle flagellar segments only very slightly transverse, seventh flagellar segment 
more transverse and slightly asymmetric, preclava with processus narrow but only slightly 
longer than body of the segment (Text-fig. 8). 

Thorax dorsally (including propodeum) 1*7-1*8 times as long as broad, densely pubescent 
on pronotum and mesonotum, pubescence moderately long, mainly dark, but sometimes whitish 
on mid lobe of mesoscutum just in front of scutellum and in front of inner margins of lateral 
lobes. Sculpture mainly rugose-reticulate, more regularly reticulate on the disc. Mesoscutum 
i*8 times to twice as broad as long. Propodeum in middle 0*43-0*45 the length of scutellum, 
with strong median carina which is about three times as long as the smooth convex dorsellum 
and only 0-35-0*38 the distance between inner edges of spiracles. Median part of propodeum, 
apart from the crenulate depressions, nearly smooth; groove formed by the flange posteriorly 
laterad of nucha in dorsal view not broader than the basal crenulate furrow. Metapleura 
rather densely hairy. Reticulate lower mesepimeron smaller than the nearly smooth upper 
part, the broad dividing depression in form of arched fovea reaching metapleural margin. 
Fore wing (Text-fig. 6) about 2 -5 times as long as broad, hyaline in basal third and on the large 
spot below marginal vein; apex faintly infumate; otherwise with two brown cross-bands, one 
appended at parastigma, the other below postmarginal vein, the bands united on disc below 
white macula. Basal cell sparsely hairy below submarginal vein. Relative lengths in holo- 
type; subcostal cell 72, marginal vein 34, postmarginal vein 34, stigmal vein 22, in two para- 
types these measures 65 : 35 : 32 : 24 and 72 : 37 : 39 : 24, resp. 

Gaster (Text-fig. 5) 1*8-1*97 times as long as broad and 1*23-1*27 times as broad as the 
mesoscutum, only 0*9-1*16 times as long as head plus thorax combined. Reticulation on 
basal halves of third and fourth tergite and on most of fifth tergite shallow, meshes mostly in 
form of transversely lengthened depressions; smooth belts at hind margins of tergites 5 and 6 
with scattered raised piliferous punctures. Posterior margins of first, second and sixth tergites 
slightly produced, arcuate; fifth tergite 2-2*24 times as broad as long, in the middle always 
distinctly shorter than tergites 3 and 4 taken together. Hairs of gaster anteriorly whitish, 
posteriorly dark. 

o*. Mainly black, with faint dark violaceous or dark green reflections on head and thorax, 
more brightly violaceous on propodeum, metapleurae and base of gaster. Antennae black; 
legs darker than in female, tibiae and tarsi blackish, also femora dorsally infuscate and usually 



EUROPEAN PTEROMALIDAE 273 

with a violaceous sheen. Fore wing slightly infumate, mostly with suffused brown clouds 
below parastigma and around stigmal vein. Head and thorax much as in female but sculpture 
relatively coarser, though in some places more superficial. The alutaceous meshes on frons 
deeper, frons therefore duller. Scrobes shallow but conspicuous. Antennae stouter than in 
female and C. laticornis; scapus three times as broad as long; pedicellus plus flagellum about 
twice as long as width of frons (49 : 25); pedicellus about 1-5 times as long as broad dorsally; 
first flagellar segment more anellus-like than in female, shorter and narrower than the second, 
which also has more adpressed pubescence like the rest of flagellum but unlike pedicellus and 
anellus; base of flagellum distinctly expanding below (in side view), the third segment the 
broadest, slightly asymmetric, slightly broader than the otherwise stoutly filiform flagellum; 
all funicle segments transverse, mostly about 1 -5 times as broad as long, with the usual thick 
semi-erect pubescence. Fore wing: relative lengths of marginal, postmarginal and stigmal 
vein as 25 : 25 : 17 (allotype; in the other paratypes as 25 : 28 : 18 and 25 : 28 : 19, resp.). 
Gaster more densely hairy than in female, hardly narrower and slightly shorter than thorax, 
with petiole distinct, though fairly transverse, smooth and convex dorsally; fifth tergite about 
as long as two preceding tergites together. Length of body 3-3-2 mm. 

Variation. Compared with the body the size of the eyes in females seems to 
vary considerably. Whilst in the holotype, breadth of the eye equals the distance 
between eyes, in one paratype of 37 mm body length, vertex breadth is 1-2 times 
the eye breadth, and in another paratype of only 3 mm length (a dwarf, reared in 
laboratory; wings still in pupal skin) the relation is 1-44 : I. In the three males 
this relation varies between 1-14 and 1-18 to 1. 

Biology. Parasite of xylophagous beetles; reared from Hylesinus toranio Bern. 
(Col., Scolytidae). 

Holotype $. Italy: Toscana, Sesto Fiorentino, vii.1943 (L. Ceresa), 'Cleonymus 
depressus (F.) det. L. Masi'; now in BMNH. 

Paratypes. Czechoslovakia: Slovakia, Zadiel, ex Hylesinus toranio, 1 $, 1954 
(A. Pfeffer); in Boucek Collection. France: Vienne, Isere, 3 <J (one of them allo- 
type) (L. Falcoz) ; in MHN, Geneva, along with 1 $ without data from Coll. Chevrier. 
Yugoslavia: Croatia, Krapina, 1 $ (Hensch); in ZIPF, Zagreb. 

Before Graham's monograph this species was frequently mistaken for Cleo- 
nymus obscurus Walker and the Czechoslovak specimen was also recorded under 
that name by Boucek (1958 : 369). 

Cleonymus laticornis Walker 

Ichneumon depressus Fabricius, 1798 : 231. Type $, France: Paris (MNHN, Paris; or lost). 

[Nee Gmelin, 1790.] 
Cleonymus laticornis Walker, 1837 : 351. Holotype <£, Ireland: Bexley (NM, Dublin). 
^.Cleonymus obscurus Walker, 1837 : 352. Lectotype (J, Britain: London (BMNH) [examined]. 

I have examined almost all the material of these two forms which Dr Graham had 
at his disposal and, thanks to several colleagues, extensive additional material of 
these rather rare insects. Special attention was paid to the variation and it was 
found that the relative length of malar space and the eye does not yield any reliable 
difference between C. laticornis and C. obscurus (used by Graham, 1969). In 33 



274 z - BOUCEK 

females with body length ranging between 3-1-57 mm, the ratio between malar 
space and eye length varied mainly between 0-5 and 0-57 (in 27 $$), not suggesting 
any gap or two-peak curve, but with two extreme deviations of 0-44 and 0-46, and 
another two of 0-65 and 0-66. The small figures do not correlate completely with 
the darker body colour, as assumed by Graham. In general, the figures obtained 
seem to suggest a trend in the eyes being relatively smaller in bigger specimens, 
which contrasts with what I found in Cleonymus brevis sp. n. The colour difference 
is notoriously unreliable with Pteromalid parasites of xylophagous beetles, but 
because the material of males, particularly of the darker form with hardly any wing 
markings (obscurus), is very scarce, I am leaving the question unresolved, although 
the study of the females suggests that only one species is involved. 
For other references and information see Graham (1969 : 38, 39). 

Material examined includes specimens from Britain, France, Switzerland, 
Czechoslovakia, Yugoslavia, Italy and Morocco (Tangier). 

EUNOTINAE 
E UNOT US Walker 

Eunotus Walker, 1834 : 297. Type-species: Eunotus cretaceus Walker, by monotypy. 

Tridymus subgen. Tritypus Ratzeburg, 1852 : 227. Type-species: Tridymus (Tritypus) areolatus 
Ratzeburg, by monotypy. 

Megapelte Forster, 1856 : 63, 66 [replacement name for Eunotus Walker, supposedly pre- 
occupied]. 

Eunotus subgen. Eunotellus Masi, 1931 : 423. Type-species: Eunotus (Eunotellus) aquisgran- 
ensis Masi; designated by Graham, 1969. 

For other references see Graham (1969). 

The genus was divided by Masi (1931) in two subgenera, Eunotus s. str., with 
5-segmented funicle in females, and Eunotellus Masi, with 4-segmented funicle in 
females. The males, however, do not yield any character in support and are 
difficult to separate even on specific level. Now another group has emerged, with 
E. hofferi sp. n. and E. kocoureki sp. n., distinguished by the relatively more flattened 
body with coarser and shallower sculpture, 3-toothed mandibles (Text-fig. 15) and 
different form of antennae in the males. Therefore, at least for the time being, it 
seems to me more appropriate to adopt species-groups rather than subgenera. 

Key to European Species 
Females 

1 Funicle 4-segmented. 

Flagellum clavate; thoracic reticulation dense; scutellum transverse, 
1-13-1-3 times as broad as long; fore wing pubescence very dense; marginal 
vein 1-34-1-63 times as long as the stigmal; first tergite smooth. 

parvulus Masi (p. 287) 

— Funicle 5-segmented ........... 2 

2 (1) First tergite distinctly reticulate, except near edges ..... 3 

- First tergite smooth ........... 5 



EUROPEAN PTEROMALIDAE 275 

3 (2) Brachypterous (Text-fig. 25) ; thorax very flat dorsally, with wide-meshed 

reticulation, meshes with smooth bottom; scutellum nearly 1-4 times as 
broad as long; propodeum very short, only one-sixth the length of scutellum; 
antenna slender (Text-fig. 10), first funicle segment transverse, the second 
subquadrate, clava about 2-5 times as long as broad . . hofferi sp. n. (p. 277) 

Macropterous ; the other characters not all present in combination ... 4 

4 (3) Scutellum distinctly though weakly convex, nearly as long as broad, very densely 

reticulate, dull; apex of scutellum bluntly angulate and reaching slightly 
beyond line with hind corners of propodeum; the latter medially extremely 
short. Sensilla linearia on flagellum very distinct; clava about three times as 
long as broad and usually darker than the mostly testaceous funicle. Marginal 
vein of fore wing 1 -5-1 -85 times as long as the stigmal, the latter bent, slightly 
longer than postmarginal vein and angle between them rather sharp, about 
30°; apex of stigmal vein about half the length of this vein from front margin 
of wing. Eye round. Body bluish black . . cretaceus Walker (p. 281) 

Scutellum (Text-fig. 13) flat, almost 1-5 times as broad as long, rather coarsely 
reticulate, fairly shiny, its apex broadly rounded and not reaching the level of 
hind corners of propodeum; propodeum in the middle at least one-third as long 
as scutellum. Sensilla of flagellum indistinct, clava hardly more than twice 
as long as broad, its first segment transverse. Marginal vein fully 25 times 
as long as the stigmal, angle between short postmarginal vein and stigmal vein 
about 45°; apex of stigmal vein about two-thirds its length from front margin 
of wing. Eye longer than broad as 18 : 16. Body greenish black. 

kocoureki sp. n (p. 279) 

5 (z) Only first flagellar segment anellus-like, the second much bigger and subequal to 

third segment. Scutellum at least slightly (1-07-1 15 : 1) longer than broad 
(steep axillulae, if seen dorsally, not included), in the apical third its sides 
converging at about a right angle ........ 6 

Basal two segments of flagellum anellus-like (Text-fig. 24), i.e., the second 
distinctly smaller than the third. Scutellum shorter than broad, usually 
broadly rounded apically ......... 7 

6 (5) Flagellum stout but hardly clavate (Text-fig. 18), its first segment narrower and 

hardly half as long as the second, which is almost as broad as the fifth; clava 
fully twice as long as broad, not distinctly asymmetric. Marginal vein less 
than twice (1-45-1 -85 times) the length of stigmal vein (Text-fig. 17). POL 
to OOL as about 2-8 : 1 ; occipital ridge often blunt, at least laterad of posterior 
ocelli. Body often about 2 mm . . . areolatus (Ratzeburg) (p. 282) 

- Flagellum strongly clavate, gradually broadening towards clava, the latter 

hardly 1-5 times as long as broad, asymmetric, subtrunctate at apex. 
Marginal vein clearly more than twice as long as the stigmal (Text-fig. 20). 
POL to OOL about 4:1; occipital ridge very sharp, even laterally. Body 
at most i-8 mm ....... obscurus Masi (p. 284) 

7 (5) Thoracic dorsum dull, densely ruguloso-reticulate. Antennal clava long-ovate, 

moderately stout, not distinctly asymmetric (Text-fig. 22). 

acutus Kurdjumov (p. 285) 
Thoracic dorsum moderately shiny, as the sculpture is more superficial. 
Antenna more strongly clavate (Text-fig. 24), often asymmetric, subtruncate 
ventro-apically ........... 8 

8 (7) Angle between stigmal and postmarginal vein very sharp, usually about 30 ; 

space between the two veins at least twice as long as broad, postmarginal 
vein at least 2/3 the marginal vein. Antennal clava fuscous, distal funicle 
segments yellowish. OOL 1-3-1-5 the diameter of lateral ocellus. 1-2- 
1 -4 mm ........ nigriclavis (Forster) (p. 286) 



276 Z. BOUCEK 

Angle between stigmal and postmarginal vein broad, about 6o°; space between 
the two veins about as long as broad, postmarginal vein half the marginal. 
Antennal clava and funicle unicolorous, yellow. OOL subequal to maximum 
diameter of lateral ocellus, i-i mm .... merceti Masi (p. 286) 

Males 

1 Antennae relatively long, flagellum plus pedicellus combined at least 1 -2 times the 

width of head; first funicle segment at least slightly shorter than the fourth, 
the latter aways distinctly elongate, in length subequal to the first claval 
segment (Text-figs 11, 14); third claval segment shorter than the first; 
flagellum with strong and high sensillar ridges, these uninterrupted on the 
whole length of the segment, 4-5 of them visible in any lateral view. Body 
rather flattened; reticulation on thorax wide-meshed; median carina of 
propodeum not distinctly raised anteriorly. Mandibles 3-toothed (Text- 
fig- 15) 2 

Antennae relatively shorter, flagellum plus pedicellus combined at most 11 
times the head width; first funicle segment usually longer than the fourth 
which, if elongate, is distinctly longer than the first claval segment (Text-fig. 
16); the latter much shorter than the third claval segment which bears two 
rows of sensilla; flagellar longitudinal sensilla much finer and shorter, but 
more numerous, forming at least on some of the basal segments two or three 
irregular rows, i.e., they are not united to form strong regular ridges along the 
whole segment as above. Body not unusually flattened; reticulation on 
thorax usually dense; median carina of propodeum raised anteriorly to form 
a distinct tooth. Mandibles 2 -toothed. ....... 3 

2 (1) Body length 0-6-07 mm; basal tergite of gaster smooth; venation of fore wing 

pale brown ......... hofferi sp. n. (p. 277) 

Body 1 0-1 -3 mm ; first tergite on disc extensively coarsely engraved-reticulate 

(as Text-fig. 13); venation of fore wing dark brown . kocoureki sp. n. (p. 279) 

3 (1) First tergite mainly longitudinally reticulate, smooth only near the edges; tibiae 

extensively testaceous; body 0-85-1-2 mm . . cretaceus Walker (p. 281) 

- First tergite smooth all over; tibae and length otherwise .... 4 

4 (3) Scutellum distinctly longer than broad (measured between axillular furrows), 

its margins posteriorly converging at about 90 ° or a slightly sharper angle. 
Inner angle of axilla 6o° or more. Postmarginal vein usually longer than the 
stigmal. Body usually at least 1 -2 mm in length ..... 5 

- Scutellum shorter, at most as long as broad but usually slightly transverse, 

posteriorly rounded or, if sides converging, the angle is more than 90°. Inner 
angle of axilla distinctly less than 6o°. Postmarginal vein often shorter than 
or as long as the stigmal. Body often less than 1 -2 mm .... 6 

5 (4) Marginal vein about 2-5 times as long as the stigmal. Body small. (According 

to Masi, 193 1 ) ....... obscurus Masi (p. 284) 

- Marginal vein about 1 -7-1 -8 times as long as the stigmal. Body often relatively 

larger, up to 2 mm, but small specimens of 1 mm are also known. 

areolatus (Ratzeburg) (p. 282) 

6 (4) Angle between postmarginal and stigmal vein very small, about 30 °; stigmal 

vein often only hardly shorter than the marginal. Body 0-75-0-95 mm. 
Flagellum brownish, combined with pedicellus often shorter than width of 
head; distal funicle segments subquadrate, constrictions between segments 
very conspicuous; pedicellus not much narrower than and distinctly more 
than half as long as the first funicle segment . nigriclavis (Forster) (p. 286) 



EUROPEAN PTEROMALIDAE 277 

- Angle between postmarginal and stigmal vein broader, usually about 45 ° or 

more; stigmal vein distinctly shorter than the marginal. Body longer than 
0-85 mm. Flagellum mostly blackish, its length with pedicellus usually 
distinctly more than width of head; funicle more compact, constrictions 
between segments usually hardly conspicuous, distal funicle segments 
elongate, the first distinctly broader and usually about twice as long as 
pedicellus ............ 7 

7 (6) Eye relatively larger (Text-fig. 26), 1. 02-1. 16 times as high as broad, maximum 
height 1 -2-i 4 times the malar space. Mid and hind tarsi, sometimes also 
tibiae, extensively testaceous. Wing pubescence very dense. 

parvulus Masi (p. 287) 

- Eye relatively smaller (Text-fig. 23), about as high as broad or slightly broader, 

its maximum height about 11 times the malar space. Mid and hind tarsi 
testaceous only basally, tibiae mostly fuscous. Wing pubescence generally 
less dense than above ..... acutus Kurdjumov (p. 285) 

Note: The male of E. merceti Masi is not known. 



The KOCOUREKI-Grovp 

Eunotus hofferi sp. n. 

(Text-figs 10-12, 25) 

$. Body black, in places with a faint dark green or bluish green tint. Antennae mainly 
testaceous but scapus infuscate except for narrowly pale apex; pedicellus also infuscate. Legs 
concolorous with body, tarsi testaceous except apical segment. Wing rudiments infumate. 
Length 11 mm. 

Head (Text-figs 12, 25) nearly 1-2 times as broad as mesoscutum, rather stout, only 1-9 
times as broad as long (in dorsal view along bottom of scrobes). POL about 6 times the OOL, 
which is hardly more than one ocellus diameter; median ocellus distinctly in front of line 
through front edges of lateral ocelli, the ocellar triangle 2-6 times as broad as high. Upper 
frons 0-43 breadth of head, rather shiny, engraved-reticulate, almost every mesh of reticulation 
with a small excentric piliferous puncture (Text-fig. 12). Lower frons with reticulation denser 
and less regular, distinctly raised on fairly concave scrobes. Eye very large, slightly higher 
than broad (16 : 14-5), without conspicuous pubescence. Other relative measurements: width 
of head 31-5, height 23, width of frons 13-7, malar space hardly 10, width of mouth about 9 
(broadening posteriorly), length of scapus 11-3, flagellum plus pedicellus 21-5, i.e., about 
two-thirds width of head. Scapus weakly sinuate, feebly thickened in basal half; pedicellus 
dorsally about 1 -5 times as long as broad, first flagellar segment anelliform (Text-fig. 10), 
narrower than the second, about o-6 times as long as broad; second flagellar segment hardly 
narrower than the third but slightly transverse, the following three segments slightly increasing 
in width and all subquadrate; clava bluntly lanceolate, distinctly broader than preceding 
segment, about as long as the four preceding segments combined. Flagellum moderately 
clavate (in terms within the genus), with sparse longitudinal sensilla which are as long as the 
segments but absent from ventral side of clava (as usual in the genus; this has caused some 
authors to regard clava sometimes as indistinctly segmented when examined from ventral 
side) . 

Thorax flattened above, slightly shorter than long dorsally. Pronotum medially about 
three-fifths the length of mesoscutum, its posterior half nearly smooth. Mid lobe of mesoscutum 
nearly 2-5 times as broad as long, notauli strongly diverging forward; mid lobe as well as 
scutellum with broad-meshed shallow, but raised, reticulation. Scutellum lateroposteriorly 



278 z. bouCek 

more finely reticulate-punctulate, itself 1-5 times as broad as long; axillar furrows strongly 
converging but reaching mesoscutum well outside of notauli. Metanotum indistinct. Propo- 
deum extremely short, about one-sixth the length of scutellum, costula not distinct, median 
carina very short but raised. Pleural parts of thorax reticulate. Legs relatively strong, 
femora moderately thickened. Wings rudimentary; fore wing triangular, obliquely truncate, 
reaching just over base of gaster (Text-fig. 25). 

Gaster slightly shorter than head plus thorax combined, strongly convex. First tergite 
about 1 -3 times as broad as long, distinctly longitudinally engraved-reticulate, sculpture fine 
and becoming obliterated towards the smooth hind margin; the latter hardly produced; 
epipleurae of first tergite with the same sculpture as its dorsum. 




Figs 10-16. Eunotus. 10-12. E. hofferi. 10, $ antenna; 11, <J antenna; 12, head with 
sculpture on frontovertex. 13-15, E. kocoureki. 13, <J, infumation of fore wing and 
sculpture of head not indicated; 14, <J antenna; 15, mandibles. 16, E. acntus, <J antenna. 



EUROPEAN PTEROMALIDAE 279 

<$. Body colour as in female. Antennae and legs brownish black, tarsi paler brown towards 
base. Wings fully developed, subhyaline. Length about 0-7 mm. Head fully 1-2 times as 
broad as mesoscutum. Relative measurements: width of head 25-5, length 12, height 19, 
irons width 14, eye 10 : 10, malar space 9, scapus 10, flagellum plus pedicellus 32. Pedicellus 
dorsally slightly elongate, about three-fifths of first funicle segment. All flagellar segments 
elongate (Text-fig. n), the first slightly shorter than the second or fourth, the latter subequal 
to first claval segment; third claval segment the smallest, narrow, subacuminate. Flagellum 
with coarse ridge-like sensilla which extend beyond apex of the segments as distinct teeth, in 
lateral view about 5 ridges visible on each segment. Thorax as usual in alate forms of genus, 
1-25 times as long as mesoscutum broad, dorsally more convex than in female, fairly shiny due 
to wide-meshed raised, but shallow, reticulation. Pronotum rather convex, its hind margin 
broadly emarginate, therefore in middle only half as long as mesoscutum; mid lobe of the 
latter about 1-7 times as broad as long. Scutellum 095 times as long as broad. Propodeum 
moderately sloping, in middle about 0-4 the length of scutellum and here distinctly produced 
beyond sublateral parts; costula irregular, less distinct than the weak median carina and the 
indicated plicae. Fore wing rather regularly and fairly densely pubescent. Relative measure- 
ments: wing length 50, width 22-5, costal cell length 17, marginal vein 8, postmarginal 3-5, 
stigmal vein 3-2 (in another specimen last three figures 7:3:3). First tergite 1-3 times as 
broad as long, dorsally smooth, but epipleurae (ventral sides of tergites) with some wide-meshed 
alutaceous reticulation as in female, but weaker. 

Apart from the antennal characters and the 3-toothed mandibles, which suggest 
close relationship with E. kocoureki sp. n., E. hofferi differs from E. parvuhis Masi 
and E. acutus Kurdjumov by its much shinier thorax. 

Biology. Host not known; all specimens collected on xerothermic grassland 
slopes, on limestone, sand or loess. 

Holotype $. Czechoslovakia: S. Moravia, Dolni Vestonice, 4.vii.io,52 (Hoffer); 
in BMNH. 

Paratypes. Czechoslovakia: Bohemia, Praha-Chuchle, 1 <$, allotype, 11.vii.1955 
(Boucek); S. Moravia, Dolni Vestonice, 1 <$, 24. v. 1954 (Hoffer); SE. Slovakia, Pilis 
Hill nr Slov. Nove" Mesto, 2 $, 3. and 13.vii.1950 (Hoffer); Kralovsky Chl'mec, 1 o*, 
20. v. 1958 (Boucek) ; paratypes partly in NM, Prague, partly in Boucek Collection. 

The species is named in honour of Dr A. Hoffer, of Prague, a prominent Czech 
hymenopterist. 



Eunotus kocoureki sp. n. 

(Text-figs 13-15) 

$. Black, with faint dark green tint mainly on head and thorax. Antennae dark testaceous, 
with pedicel infuscate and scapes also slightly infuscate in basal halves. Legs concolorous 
with body, fore basitarsus basally and mid and hind tarsi except the claw segment dark 
testaceous. Fore wing infumate but paler basally at hind margin and in apical third ; venation 
dark brown. Length 1-4 mm. 

Head broader than mesoscutum as 41 : 35, in dorsal view 2-2 times as broad as long, strongly 
crescentic, with occiput strongly excavated, but dorsal ridge not sharp. POL to OOL about as 
4:1, OOL itself equals about 1 -4 diameter of ocellus; line drawn through front edges of lateral 
ocelli intersecting hind quarter of median ocellus, ocellar triangle about 3-8 times as broad as 



280 Z. BOUCEK 

high. Upper frons in front of ocelli nearly half (0-47) as broad as head, rather dull, with 
conspicuous engraved reticulation, each mesh bearing a piliferous puncture which takes up at 
least half of mesh surface. Lower frons with very dense raised reticulation, this denser in the 
middle, the very shallow scrobes therefore still duller; sublaterally reticulation less dense and 
less regular, at orbits and on genae again very dense. Relative measurements: head width 41, 
height 27, frons width 19-5, eye 15 : 17-5, malar space 13, mouth width about 11. Eye 
pubescence sparse, short, inconspicuous. Both mandibles 3-toothed. Relative length of 
scape 15, flagellum plus pedicellus 27, i.e., slightly less than width of head less one eye. Scapus 
slender, distinctly sinuate, scarcely thicker in basal half; pedicellus dorsally nearly twice as 
long as broad (Text-fig. 13) ; flagellum slender, weakly clavate, its first segment shorter but not 
abruptly narrower than the second and still slightly longer than broad ; longitudinal sensilla of 
flagellum sparse, long, 1-2 in a view on second and third segment and 3 on fourth and fifth 
segment; clava hardly longer than three preceding segments combined, long-oval, nearly three 
times as long as broad, with three rows of sparse sensilla, its first segment slightly longer than 
the preclaval one and distinctly longer than the third claval segment. 

Thorax dorsally flattened, broad (Text-fig. 13), from anterior edge of collar down to apex of 
propodeum only as long as breadth of mesoscutum. Pronotum dorsally 0-45 the length of 
mesoscutum, laterally narrowed, without shoulders; front half of collar finely rugulose-reticulate, 
hind half nearly smooth, shiny ; lateral panel of pronotum above with deep depression delimited 
by a horizontal crest opposite to lower edge of the small prepectus. Mid lobe of mesoscutum 
twice as broad as long, together with scutellum coarsely and rather deeply reticulate-punctured; 
side lobes and axillae finely sculptured, partly smooth at cross-suture. Scutellum 1 -45 times as 
broad as long, apically broadly rounded, apical quarter much more finely and superficially 
sculptured than the disc; axillar furrow reaching mesoscutum just outside notaulices; axillulae 
not distinct. Metanotum visible only laterally. Propodeum subhorizontal, medially slightly 
more than one-third the length of scutellum; median carina low, rather broad, weakly raised 
before middle; costula rather indistinct, crossing the mainly longitudinal rugae; spiracle oval, 
open; hairs of callus short, not conspicuous. Thoracic pleurae dull, rather deeply reticulate, 
except below the wings where sculpture forms longitudinal striae ; mesepimeron without central 
pit; metapleura and lower part of lateral panel of propodeum slightly shiny. Fore wing fully 
developed, about 2-6 times as long as broad; its hind margin with distinct lobe beyond basal 
quarter; front margin shallowly emarginate at end of costal cell. Relative measurements: 
length of costal cell 27, marginal vein 10, postmarginal vein 3-5, stigmal vein 4. Wing disc 
with dense but extremely short hairs, in basal two-fifths hairs much longer and rather sparse. 

Gaster with slightly protruding ovipositor sheaths as long as head plus thorax combined 
(Text-fig. 13). First tergite 1-3 times as broad as long, its hind margin produced slightly 
arcuately, dorsal and epipleural surface distinctly longitudinally striate-reticulate, but smooth 
basally and near edges, more broadly so in hind corners. 

o*. Differs from female as follows. Antennae mostly black, scapus sometimes dark testaceous 
at both ends or only distally. Apex of mid and hind tibiae shortly testaceous. Fore wing 
subhyaline, mostly only with slight infumation forming an angulate streak below parastigma 
along basal and cubital folds. Length 1 -o-i -2 mm. Head more regularly reticulate-punctured, 
frons above duller, reticulation raised. In dorsal view temples visible though strongly receding. 
Eyes relatively smaller; their relative measures 12 : 13, width of frons 19-5, malar space 12, 
mouth width 10, scapus 13 (nearly as slender and sinuate as in female), flagellum plus pedicellus 
combined 44, i.e., four-thirds the head width. Pedicellus subglobular, about half as long as 
first funicle segment. All flagellar segments (Text-fig. 14) with high coarse sensillar ridges 
along the whole lengths which project like teeth beyond apex of each segment; groove-like 
bottoms slightly shiny, with short regular semi-erect hairs. Flagellum slightly tapering to- 
wards apex, fourth segment about 1 -7 times as long as broad, slightly longer than the first but 
hardly longer than the following first claval segment; third claval segment the smallest, dis- 
tinctly shorter than any other flagellar segment. Scutellum regularly reticulate-punctured all 
over. Gaster hardly half as long as head plus thorax together, first tergite distinctly longi- 
tudinally engraved-reticulate. 



EUROPEAN PTEROMALIDAE 281 

Biology unknown. Collected on xerothermic grassland. 

Holotype $. Bulgaria: Sandanski, vi.ig69 (Kocourek); in BMNH. 

Paratypes. Bulgaria: Sandanski, 4^ (including allotype) (Kocourek); Boucek 
Collection. 

Named in honour of Ing. M. Kocourek, a Czech hymenopterist and very good 
collector. 

Eunotus hofferi and E. kocoureki form a very distinctive species-group. Unlike the 
other species of the genus they have 3-toothed mandibles (Text-fig. 15), in males the 
longitudinal sensilla of the flagellum are very strong, ridge-like and extend along the 
whole segment and the segments of the funicle and clava are subequal, as stressed in 
the key. The females, apart from the rather depressed thorax with the scutellum 
very short and transverse, have a peculiar depression on the upper part of the lateral 
panel of the pronotum ; this fovea is present though less conspicuous, in the males. 
Both species seem to be associated with xerothermic habitats although nothing is 
yet known about their hosts. 



The CRETACEUS-group 

Eunotus cretaceus Walker, although rather distinctive, seems to be more related 
to E. areolatus and E. obscurus Masi than to the above group or the aa^ns-group. 

Eunotus cretaceus Walker 

Eunotus cretaceus Walker, 1834 : 298. Lectotype $, Britain: Isle of Wight (BMNH) [ex- 
amined], 

Eunotus festucae Masi, 1928 : 128. Syntypes <J<J, $$, Italy: San Vito near Modena, ex 
Eriopeltis festucae (Fonscolombe) (MCSN, Genoa and BMNH) [examined]. 

For comments on the type-material and synonymy see Masi (1931) and Graham 
(1969). I have seen Masi's material in Genoa but at that time (1965) I did not 
select the lectotype. 

Biology. E. cretaceus develops as predator on the eggs (probably all species of 
the genus develop in a similar way) of the following Coccids on grasses: Eriopeltis 
festucae (Fonscolombe) (e.g., Masi, 1928 and 1931), E. agropyri Borchsenius (n. rec), 
E. Istrelkovi Borchsenius (Graham, 1969), Eriopeltis sp. (n. rec), Scythia (=Mohelnia) 
festuceti (Sulc) (n. rec). 

Distribution. Europe, from Britain and southern Sweden to southern Italy 
and to the U.S.S.R. (Moldavian S.S.R. and Georgia). 

Material examined. 
Type-data given in synonymy. 

B 



282 Z. BOUCEK 

Britain: Surrey, White Downs near Dorking, i $, 5^.1964, 1 <$, 20. vi. 1970 
(Boucek) ; Warwickshire, Stratford-on-Avon, ex Eriopeltis festucae, 4 9, 4 <$, i6.viii. 
1965 (R. E. Evans), BMNH. Sweden: Skane, Kivik, ex E. festucae, 4 9, 1 g, viii. 
1938, and Loderup, 1 $, 22.vii.1938 (D. M. S. & J. F. Perkins), BMNH. Germany: 
'Deutschland', 1 $ (Erichson), MNHU, Berlin, identified as E. festucae by Masi. 
Czechoslovakia: Bohemia, Mila, Stfedohofi Mts, 1 S, 5.VU.1956 (Boucek); Praha- 
Hanspaulka, ex Eriopeltis festucae, 14 9, 3 <$, about 1945 (B. Stary) ; Praha-Kosife, 
1 J, i.vi.1947 (Boucek) ; Mala Skala near Turnov, 1 9, 1942 (Obenberger) , NM, Prague; 
Nove Mesto nad Metuji, 1 9. vii.1961 (/. Macek); Moravia, Mohelno, 2 9. 6.VH.1957 
(Boucek) ; Slovakia, Kovacov near Sturovo, ex Scythia festuceti, 6 2, 1 d, i960 
(Hoffer); Mt Revan, 1200 m, 1 9, 5.ix.i956 (/. Brtek). U.S.S.R., Moldavian 
S.S.R.: Slobodzeia, ex Eriopeltis agropyri, 4 9. viii. i960 (Talitzki). Georgia: 
Tbilisi district, ex Eriopeltis sp., 1 9> 3 S> v i- IO <57 (Hoffer). 



Eunotus areolatus (Ratzeburg) 
(Text-figs 17-19) 

Tridymus {Tritypus) areolatus Ratzeburg, 1852 : 227, 2 figs. Type $, W. Germany: Hohen- 

heim [lost]. 
[Enargopelte obscurus (!) Forster; Kryger, 1943 : 79-81, figs 5, 6. Misidentification.] 
Eunotus subcyaneus Erdos, 1953 : 222-223, fig- x - Holotype $, Hungary: Kelebia ^TM, 

Budapest) [examined]. Syn. n. 
? Eunotus antshar Nikolskaya in Nikolskaya & Kyao, 1954 : 4 I 3 _ 4 I 4' fig s 5 a_ d- LECTO- 
TYPE $, U.S.S.R.: W. Kazakhstan, Yanvartsevo (ZI, Leningrad), here designated [exam- 
ined]. 

This is the only species known to me which has the scutellum elongate and, in the 
female, the antenna with one anellus only and the flagellum rather stout, not much 
attenuate towards the base. Both these characters are seen in Ratzeburg's figure. 
This author also described the propodeum as having a median areola (on the basis of 
which he erected the subgenus Tritypus). As a matter of fact the areola is not 
developed in any Eunotus. But in the present species the median part of the 
propodeum protrudes posteriorly more than in any other species and being separated 
from the sublateral parts by a distinct costula, it may, in some lights and at lower 
magnification, and when seen slightly from behind, suggest a presence of a median 
area (Text-fig. 19). In such a view the median carina often becomes obsolete, while 
the costula is distinct, in middle angulate forward. E. areolatus is also the largest 
known species in Europe and this again fits Ratzeburg's statement of 2". He 
received his specimens from a 'Coccus' on Salix aurita in south-western Germany. 
This suggests a host not associated with warm places and indeed, most material at 
hand comes from moister and colder habitats. 

I have seen also Kryger's material from Denmark kindly made available by 
Mr Bakkendorf and the type of E. subcyaneus Erdos on which the first vital informa- 
tion was supplied by Prof. Szelenyi, who later kindly submitted the type itself. I 
have little doubt that E. antshar also is the same. Its lectotype female, kindly 



EUROPEAN PTEROMALIDAE 



283 



submitted by Dr Trjapitzin, differs from the two females of areolatus presently at my 
disposal in having the antennal scape slightly stouter (3-4 : i), the malar space 
slightly shorter than eye (25 : 28; in areolatus from Sweden 30 : 29 and 25 : 25, 
resp.), and the propodeal basal tooth in lateral view slender and jutting out towards 
the scutellum. These differences are small but may prove eventually to be constant, 
when more is known. 




Figs 17-26. Eunotus. 17-19. E. areolatus. 17, fore wing venation in §; 18, $ antenna; 
19, thorax. 20-21. E. obscurus. 20, fore wing venation in $; 21, $ antenna. 22-23. 
E. acutus. 22, 9 antenna; 23, <J head in facial view. 24, E. nigriclavis, $ antenna. 
25, E. hofferi, body of 9. 26, E. parvulus, head of <$ in facial view. 



284 Z. BOUCEK 

Biology. Ratzeburg (1852) obtained his E. areolatus from a 'Coccus' on Salix 
aurita, Kryger (1943) collected it on Salix repens, Erdos (1953) on Pinus nigra. 
Eunotus antshar was reared from Rhodococcus spiraeae (Borchsenius) (Nikolskaya & 
Kyao, 1954). The species, in common with the following E. obscurus, seems to be 
associated with Coccids on trees and bushes, not on grasses. 

Distribution. Sweden, Denmark, W. Germany, Czechoslovakia, Hungary; 
PU.S.S.R. (W. Kazakhstan). 

Material examined. 

Type-data of E. subcyaneus and of E. antshar given in synonymy. 

Sweden: Upland, Vallentuna, 1^,1$, 17.iv.1960 (K. J. Hedqvist); Skane, Hoor 
district, 1 $, n.vi.1938 {D. M. S. &J. F. Perkins), BMNH. Denmark: N. Sealand, 
Lyngby nr Copenhagen, 1 $, 9.vi.i962 (Bakkendorf) ; Sandkroen, 1 $, 26.V.1931 
{Kryger), UZM, Copenhagen. Czechoslovakia: Bohemia; Jedlova near Rumburk, 
1 cJ, 29. vi. 1957 (Boucek); Moravia, Hodice near Jihlava, 3 $, 7.VL1953 (F. Kodys). 



Eunotus obscurus Masi 

[Eunotus cretaceus Walker; Masi, 1907 : 262-266, figs 23, 24. Misidentification.] 

Eunotus obscurus Masi, 1931 : 424, 428-430, fig. ia. Syntypes $? and 1 q\ France (MHN, 

Paris), Germany (NM, Vienna), Italy: Bevagna (MCSN, Genoa) and Spain (IEE, Madrid) 

[mostly examined]. 
[Eunotus cretaceus Walker; Kryger, 1943 : 75—78, figs 3, 4. Misidentification.] 

I have seen most of the syntypes of E. obscurus and compared them with my 
specimens, but at that time (1965, 1966) did not select the lectotype. Kryger's 
material was also examined. The latter author (1943) mentions having reared 35 
females but no male. Neither have I seen any male in spite of having had more 
varied material. The only male known is the one recorded by Masi (1931) from 
Spain. Eunotus obscurus may be parthenogenetic, at least in northern and central 
Europe. Even the description of the Spanish male, judging from the unusually 
long marginal vein, may concern the closely related Eunotus areolatus (Ratzeburg). 

Biology. The parasite attacks Coccids on various bushes and trees, mainly 
Pulvinaria vitis (L.) (=betulae L.) (Masi, 1931 and n. rec; probably also 'Coccus on 
Salix repens concerns this species; Kryger, 1943). Another record is Parthenole- 
canium persicae (F.) on Robinia pseudacacia L. (Masi, 1931). 

Distribution. Denmark, W. Germany, France, Spain, Italy, Czechoslovakia, 
U.S.S.R. (Moldavian S.S.R. and ? Uzbekistan). 

Material examined. 

Type-data given in synonymy. 

Denmark: N. Sealand, Sandkroen, ex Coccus on Salix repens, 4 $, coll. vi.1929, 
em. v.1930 {Kryger). Czechoslovakia: E. Slovakia, Kosice, 1 $, 31.V.1952 
{Kocourek). U.S.S.R., Moldavian S.S.R. : Dubossary, ex Pulvinaria betulae on 



EUROPEAN PTEROMALIDAE 285 

Crataegus, 4 $, 2.VL1964 (Talitzki); Kishinev, ex Pulvinaria betulae, 8 $, i.vi.1964 
(Talitzki). U.S.S.R., PUzbekistan: Agashik, ex Pulvinaria betulae, 4 $, 9.viii.i928 
(Archangelskaja) , in ZI, Leningrad. 



The ACUTUS-group 

Eunotus acutus Kurdjumov 
(Text-figs 16, 22-23) 

Eunotus acutus Kurdjumov, 1912 : 330-331, figs iA-D, 3A-B. LECTOTYPE ?, Ukrainian 
S.S.R. : Poltava (ZI, Leningrad), here designated [examined]. 

According to a personal statement by the late M. N. Nikolskaja, all that was left 
of the Kurdjumov collection at the Experimental Station in Poltava, where he 
worked, was transferred to the Zoological Institute in Leningrad about 40 years ago. 
The only type-material of E. acutus consists of two specimens. One syntype is in 
bad condition (most of thorax only left) and labelled '13/46' ; the other one, a female 
in good condition, is selected here as lectotype. It is labelled '13/45' and, probably 
in Kurdjumov's handwriting, 'Eunotus acutiventris Kurd}.'. He probably changed 
the name in the manuscript, as may be guessed also from the apparent derivation of 
the name from the gaster which he described in the key as 'acute angled at the tip'. 
It fits the description well and agrees exactly with one of the smallest specimens 
from Bohemia. The body-size varies in females from 1—1*5 mm » m males 0-85-1-2 
mm. 

The females can be recognized rather easily on the characters given in the key 
above, but the males are very similar to those of E. parvulus Masi and, to some 
extent, also to E. nigriclavis (Forster). In the former two species I have found also 
a rather wide range of variation and the rather slight differences mentioned in the 
key above proved only more or less reliable. The figures obtained by careful 
measurement often overlap. For example, among more than 20 males of E. acutus 
(reared with the females) the ratio of the breadth of the frons and the breadth of the 
eye is 1-34-1-5 : 1, whilst in 18 males of E. parvulus (mostly British and North 
European specimens) it is 1-08-1-35 : x - Otherwise in E. parvulus the vestitute of 
the eyes seems to be generally longer, more conspicuous. I have not found any good 
character in the relative lengths of the fore wing veins, in scutellum (which seems, 
however, to be relatively broader in E. parvulus), or in propodeum, thoracic pleurae, 
etc. In some males of E. parvulus the scapus is more attenuate distally, but in 
some others it is not. In both species the thoracic dorsum is rather dull. In the 
females of E. nigriclavis it is shinier, and the scutellum is slightly more convex. 

Biology. The records from the fresh material together with information 
published earlier list the following Cocci ds as hosts: Acanthococcus greeni (Newstead) 
(Kurdjumov, 1912), Rhizococcus agropyri Borchsenius (n. rec.) and Greenisca placida 



286 Z. BOUCEK 

(Green) (n. rec). The parasite seems to be closely associated with grasses, mainly 
with Agropyrum species in xerothermic habitats. 

Distribution. Poland, Czechoslovakia, Ukrainian S.S.R. 

Material examined. 

Type-data given in synonymy. 

Poland: Poznah-Bebice, ex Rhizococcus agropyri on 'low grass', 5 $, 1 <$, vii.-xi. 
1967 (Lewandowski). Czechoslovakia: Bohemia, Hazmburk Hill near Libo- 
chovice, 1 $, 2.VL1943 (Hoffer); Praha-Ruzyne, 1 $, 22.V.1953 {Boucek); Karlstejn, 
ex Greenisca ?placida, 60 $, 34 $, 1957-58 {Hoffer) ; Prachatice, 1 $, 30.viii.1950 
{Hoffer); Moravia, Hostyn, ex Greenisca placida, 5 9, 1 <$, 1957 {P. Stary). 

Eunotus nigriclavis (Forster) 

Megapelte nigriclavis Forster, 1856 : 66. Holotype $, Germany: Aachen (NM, Vienna) 
[examined] . 

The material mentioned below has been compared with the type of E. nigriclavis 
in Vienna. The form of the antennal clava does not seem to be such a good character 
as Masi thought. Although in this and the following species the clava is usually 
stouter than in any other species, in all species it has the ventral side deprived of 
longitudinal sensilla in place of the extended area of micropilosity and, in E. nigri- 
clavis, appears usually slightly obliquely truncate in dry specimens. 

Biology. Host not known. In central Europe E. nigriclavis occurs mainly in 
woods and montane regions. 

Distribution. Germany (West), Czechoslovakia, Yugoslavia. 

Material examined. 

Type-data given in synonymy. 

Czechoslovakia: Bohemia, Krkonose Mts, Cerna Hora, 1200 m, 1 $, ix.1949 
{Hoffer) ; Novy Hradec Kralove, 250 m, 1 $, 6.viii.i958 {Boucek) ; Hfirka v Posumavi, 
1 $, 17.vii.1954 {Hoffer) ; Sumava Mts, Horni Snezna, 1000 m, 1 <$, 15.vii.1946 
{Hoffer); Moravia, Javofice, 800 m, 1 $, 7.viii.i944 {Hoffer). Yugoslavia: Dal- 
matia, Biograd na moru, 1 $, 14.vii.1968 {Boucek). 

Eunotus merceti Masi 

Eunotus merceti Masi, 1931 : 424, 433-435, fig. 2. Holotype $, Spain: El Pardo (IEE, Madrid) 
[examined]. 

Very similar to E. nigriclavis but differing mainly in the fore wing venation. 
Only one female known. 

Biology unknown. 

Distribution. Spain. 



EUROPEAN PTEROMALIDAE 287 

Species sola 
Eunotus parvulus Masi 

Eunotus (Eunotellns) parvulus Masi, 1931 : 424, 435-437, figs 3a-d, <^$. LECTOTYPE $, 

Austria: Wimpassing (NM, Vienna); here designated [examined]. 
Eunotus (Eunotellus) aquisgranensis Masi, 1931 : 424, 437-438. Holotype ?, W. Germany: 

Aachen (NM, Vienna) [examined]. Syn. n. 

Out of i $ and 4 $ of the original material of E. parvulus, 1 <$ from Bohemia and 
1 9 from Austria were designated by Masi as 'types'. Graham (1969 : 74) overlooked 
Masi's practice in designating types representing both sexes and mentions only the 
male as 'type' (and the other specimens as paratypes). I select the female as lecto- 
type, as males of this species are not always safely recognizable. 

Masi (1931) regarded E. parvulus and E. aquisgranensis as two different species, 
mainly on the difference in the relative lengths of the marginal and stigmal veins. 
I have found this character unreliable and have failed to find others. In 7 females 
examined the marginal vein is from 1 -34 to 1 -64 times as long as the stigmal vein, 
in another female (from Sweden) i-8. The males are not always safely identifiable 
and that is why I measured only the specimens from Britain, where the occurrence 
of another similar species really is minimal. In 10 males the marginal vein showed 
ratio towards the stigmal vein between i-6 and 2-1. The figures seem to be higher 
than in the females, but oddly enough among the females the highest figure, 1 -64, is 
shown again by a British specimen. A similar finding, though in one female only, 
led Graham (1969 : 74) to regard his specimen as E. aquisgranensis. 

Distribution. Britain, Sweden, Germany (West), Austria, Czechoslovakia. 

New records. Britain: Esher Common, Surrey, 3 <$, 21. vi. 1970 (Boucek); 
Chobham Common, 1 $, 2 $, 19. vi. 1970 (Boucek); Bald Hill nr Lewknor, Oxford- 
shire, 4^, 13. vi. 1970 (Boucek); Wytham Wood, Berkshire, 1 <$, 26. vi. 1964 (Boucek; 
published as Eunotus sp. in Boucek, 1965 : 83). Sweden: Vasterbotten, Norsjo, 
1 9> 5- vn - I 956 (Sundholm); Blekinge, Sjoarp, 1 $, i2.vii (Hedqvist). Czecho- 
slovakia: Bohemia, Kfesin nr Libochovice, 1 $, 31.vii.1943 (Hoffer); Praha- 
Chuchle, 1 (J, 1 9, 24.vi.1955 (Boucek); Slovakia, Slov. Nove Mesto, Pilis Hill, 1 9» 
31.V.1952 (Hoffer); Somotor, 1 9. i.vii.1952 (Kocourek). Austria: 'Vimpacs' 
(= Wimpassing), Leitha Mts, 1 9» 20.vii.1915 (Ruschka), NM, Vienna. 

MISCOGASTERINAE 
SUSTERAIA gen. n. 

(Text-figs 29-32) 
Type-species : Susteraia acerina sp. n. 

Body metallic. Head and thorax very finely raised-reticulate, pilosity inconspicuous. 
Occiput not margined, temples and genae terete. Scrobes shallow. Lower face not pro- 
tuberant. Clypeus very finely reticulate-punctulate, sub trapezoidal, weakly transverse, its 



288 Z. BOUCEK 

upper margin indicated by impressed line, diverging lateral margins slightly raised below, lower 
margin with distinct tooth in the middle, broadly emarginate on either side (Text-fig. 31). 
Malar space not concave. Mandibles with lower margin lamelliform at base, margin nearly 
straight. Labio-maxillary complex normal in both sexes. Antennae inserted near centre of 
face, well above lower ocular line, in both sexes 13-segmented, filiform, with two anelli, all 
funicle segments in female elongate, decreasing in length, the first longer than the pedicellus; 
clava with both sutures nearly perpendicular. In male antenna similar, though slightly longer, 
each funicle and clava segment with several dense irregular rows of sensilla linearia but other- 
wise nearly bare, with sparse microscopic semidistant hairs (Text-fig. 30). 

Thorax fairly convex and elongate (Text-fig. 29). Pronotum rounded, rather short medially, 
lateral panels shallowly concave. Notauli complete, broad and deep, on bottom smooth 
between crenulae, as well as the axillar furrows which meet the mesoscutum well inside of 
notauli. The scuto-scutellar suture fairly sinuate. Scutellum elongate, convex, broadly 
meeting mesoscutum, frenal groove straight, in 3/4 of scutellum, frenum more coarsely reticulate 
than the disc; axillulae distinctly separated, bearing some longitudinal rugae. Dorsellum 
convex, shallowly alutaceous. Propodeum transversely strongly convex, posteriorly deeply 
emarginate, foramen bordered by high reflexed carina which extends to smooth crescentic 
nuchal strip in the middle; median carina fine, as long as frenum; plicae and nucha absent; 
spiracles of medium size, elliptical, removed from anterior margin; part behind them deeply 
depressed down to narrow supracoxal flange; callus convex, generally with one row of weak 
hairs. Prepectus broad, reticulate, without oblique carina, groove-like along upper and hind 
edge near tegula. Mesepisternum reticulate, on ventral side with some hairs, epimeron mainly 
smooth, at least above, in middle with asymmetrically arched groove. Metacoxa reticulate, 




/ / / ' ^- 



Figs 27-32. 27-28. Semiotellus rujanensis. 27, apex of $ antenna; 28, fore wing 
venation. 29-32. Susteraia acerina. 29, body of $ in lateral view; 30, o* flagellum with 
pedicellus; 31, head of £ in facial view; 32, fore wing venation in $. 



EUROPEAN PTEROMALIDAE 289 

bare dorsally, laterally broadly depressed. Legs slender, hind tibia with two spurs. Fore 
wing with slender venation (Text-fig. 32), without hyaline break at end of parastigma; marginal 
vein about twice the stigmal and slightly shorter than the postmarginal ; stigma small; pubes- 
cence weak, not dense, marginal ciliae short, basal fold with some hairs, basal cell bare and 
open below as well as speculum which extends far below marginal vein. 

Gaster in female long and strongly compressed from sides, hypopygium reaching about 
one-third along, its tip emarginate. Petiole extremely short, in middle linear. First tergite 
short, its hind margin weakly emarginate in the middle; second tergite the shortest, the sixth 
the longest, about 1 -5 times as long as epipygium which is rather large, unusually high in lateral 
view (Text-fig. 29), cerci at its lower margin in apical fifth, their bristles short. Ovipositor 
shortly protruding. Gaster finely transversely alutaceous, on sides and at apex with in- 
conspicuous dark hairs. Male gaster narrow and short; petiole moderately transverse, smooth. 

Named in honour of my old friend and teacher, Mr O. Sustera of Prague, a 
hymenopterist, who more than 60 years ago proposed the first reasonable basis for 
the taxonomy of the European Pompilidae. 

Susteraia gen. n. belongs to Miscogasterinae, tribe Miscogasterini and in Graham's 
key to genera (1969 : 150-155) it may be easily keyed out by inserting the following 
paragraphs instead of 18 (17) : 

18 (17) Antennae short, in $ combined length of flagellum plus pedicellus less than, in 6* 
at most equal, to the breadth of head; pedicellus longer than the first funicle 
segment; lower margin of clypeus laterally excised, in the middle with a 
broad tooth accompanied on either side with another weaker tooth; inter- 
antennal space convex, much broader than diameter of torulus. 

NODISOPLATA Graham 
Antennae much longer, flagellum plus pedicellus much longer than breadth 
of head; pedicellus shorter than first funicle segment; clypeal margin 
shallowly emarginate laterally and with a simple, rather sharp tooth in the 
middle ; interantennal space not broader than torulus . . . . 18a 

18a (18) Mid lobe of mesoscutum and scutellum flat; scutellar disc delimited by 
axillular furrows and frenal furrow subquadrangular, with sides nearly 
parallel, surface delicately engraved-reticulate; gaster not long, dorsally 
flat, not compressed from sides; propodeum smooth or nearly so, median 
carina indistinct, petiolar margin low . . . KSENOPLATA Boucek 

Mid lobe of mesoscutum and scutellum very distinctly convex, sides of 
scutellum clearly converging forward, the disc densely reticulate, not shiny; 
gaster long, in $ distinctly compressed from sides; propodeum reticulate- 
rugulose, with distinct median carina .... SUSTERAIA gen. n. 

Susteraia acerina sp. n. 

(Text-figs 29-32) 

$. Body mainly vivid cupreous, head and thorax in places slightly violaceous, thoracic 
pleurae and gaster basally red to golden, gaster dorsally and posteriorly mostly dark purple. 
Scapes, pedicels, mouth parts, tegulae and legs apart from coxae, reddish testaceous; tarsi 
paler except apex. Wings hyaline, venation testaceous. Length 3-8-4-1 mm. 

Head 1-16 times as broad as mesoscutum, dorsally twice as broad as long, with temples 
hardly one-third as long as eyes, rounded, receding. POL about 2-1 times OOL. In facial 
view head about 1 -26 times as broad as high. Lower face at mouth margin with 6 longish hairs, 
which are about three times as long as the normal inconspicuous pubescence. Relative measure- 



2 go Z. BOUCEK 

ments: head width 58, height 46-6, frons width 31, eye 30-5 : 22, malar space 9, width of mouth 
27, length of scape 21, pedicellus plus flagellum 82. Pedicellus dorsally twice as long as broad, 
first funicle segment 2-1 times, the fifth 1-4 times, the sixth 11 times as long as broad; clava 
hardly longer than two preceding segments together. 

Thorax dorsally (collum not measured) about 1 -7 times as long as breadth of mesoscutum. 
Pronotum without conspicuous smooth belt at hind margin, which is thin. Mid lobe of meso- 
scutum fully as long as broad. Scutellum, if axillulae excluded, about 1 -3 times as long as broad. 
Propodeum extremely finely rugulose-reticulate, sublaterally from hind margin with a few 
longitudinal rugae; hind corner above coxa with a lobate supracoxal lamina. Fore wing relative 
measurements: length 184, width 70, subcostal cell 70 : 6, marginal vein 34, postmarginal 41, 
stigmal vein 17, distance between upper margin of stigma and postmarginal vein 9. Lower 
surface of costal cell with one row of hairs, sometimes narrowly interrupted in middle, doubled 
apically (Text-fig. 32). 

Gaster very narrow, 1-4-1-45 times as long as head plus thorax combined (Text-fig. 29). 

o*. Golden-cupreous, only gaster posteriorly dark purplish; flagellum beneath ochreous; 
scapus and legs except metallic coxae, pale testaceous. Length of body 2-8 mm. Relative 
size of eye 24 : 18, scapus 14 : 4, flagellum plus pedicellus (Text-fig. 30) 79, i.e., nearly 1-65 
times the width of head. Pedicellus dorsally 1 -2 times as long as broad, half as long as first 
funicle segment which itself is 2-5 times as long as broad, the sixth i-6 times as long as broad, 
clava as long as two preceding segments combined. Fore wing slightly broader than in female, 
134 : 57, relative length of marginal vein 27, postmarginal 29, stigmal vein 15, distance between 
stigma and postmarginal vein only 1-9 times the height of stigma. Gaster narrow, slightly 
shorter than thorax. 

Biology. After receiving the specimen reared in the Ukraine, with the suggestion 
that it might be a parasite of a weevil in maple seed, I mentioned the matter to my 
colleague Dr Strejcek, a keen coleopterist working mainly on Curculionidae. He 
collected various samples of seeds and actually succeeded in rearing one male of the 
parasite, but no species of the Curculionid genus Bradybatus which we presumed to 
be the host. It is possible, however, that the Bradybatus species leave the maple 
seed earlier, or that the damaged seed falls earlier as does the seed of Sorbus attacked 
by Megastigmus brevicaudis Ratzeburg (Hym., Torymidae). The taxonomic 
affinity of Susteraia acerina cannot exclude also a possibility that a Dipteron is the 
host, as may be suggested by a nice Torymus species reared from the same lot of 
maple seed. 

Holotype $. Czechoslovakia: Bohemia, Starkoc near Nachod, vii.1955 (J. 
Macek) ; in Boucek Collection. 

Paratypes. Czechoslovakia: Bohemia, Praha-Krc, ex seed of Acer pseudo- 
platanus, 1 $ (allotype), ii.1969 ex seed collected xii.1968 {Strejcek). Ukrainian 
S.S.R. : Kiev, Botanical Garden, ex Pweevil in seed of Acer pseudoplatanus, 1 $, 
13.iii.1968 (M. Zerova). 



SEMIOTELLUS Westwood 
(Text-figs 27, 28) 

Semiotus Walker, 1834 : 288, 290. Type-species: Semiotus mundus Walker; designated by 

Westwood, 1839. [Homonym of Semiotus Eschscholtz, 1829.] 
Semiotellus Westwood, 1839 : 70. [Replacement name for Semiotus Walker.] 



EUROPEAN PTEROMALIDAE 291 

Semiotellus rujanensis sp. n. 

(Text-figs 27, 28) 
Semiotellus sp. indet., Graham, 1969 : 254, 255, ?. 

$. Bluish green; the following parts testaceous: knees, fore tibiae, narrow apices of mid 
and hind tibiae, tarsi except at apex. Wings hyaline, venation dark brown. Length 3-4- 
3'5 mm. 

Head 11 times as broad as mesoscutum, in dorsal view about twice (in holotype 2-06 times) 
as broad as long, in facial view 1-27 times as broad as high. Piliferous punctures coarse, very 
distinct, numerous, but wanting laterad of paired ocelli, in front of median ocellus and on 
lower face dorsad and laterad of clypeus. POL 17 times OOL. Area between clypeus and 
antennae rather protuberant but not very convex transversely, separated from clypeus by 
deep furrow between the very deep large tentorial pits. Clypeus slightly transverse, minutely 
reticulate, its lower margin slightly arched, produced. Mandible 2-toothed, the upper edge of 
upper tooth broad, nearly straight, not notched. Malar space with deep fovea just behind 
upper end of malar sulcus. Relative width of frons 42, of head 65, eye 29 : 21, malar space 15, 
width of mouth 32, scape length 23, flagellum plus pedicellus 63. Scapus laterally 3-7 times as 
long as broad; pedicellus dorsally i-6 times as long as broad, slightly longer than first funicle 
segment; basal funicle segments slightly elongate, the fifth subquadrate; flagellum moderately 
clavate, the segments slightly increasing in width up to second claval segment; clava in lateral 
view (Text-fig. 27) 2-4 times as long as broad, dorso-apically subtruncate, bearing here an 
extensive slightly convex area of micropilosity ; the first claval suture perpendicular, the second 
distinctly oblique. 

Head and thorax with inconspicuous, usually short, mainly dark pubescence, in spite of 
broad and very distinct piliferous punctures; the latter sparser on disc of mid lobe of meso- 
scutum and nearly absent from posterior half of scutellum. Apex of scutellum raised in middle. 
Propodeum medially one quarter the length of scutellum, steeply elevated into median ridge 
(rather than carina), all over finely reticulate, with some rather fine irregular rugae; plicae 
distinct, straight; spiracle large, its diameter equal to breadth of the lateral smooth strip of 
metanotum. Basal cell of fore wing bare, basal and cubital folds also nearly bare, with at most 
one hair. For venation see Text-fig. 28. Relative length of marginal vein 42, postmarginal 25, 
stigmal vein 13, distance of the slightly enlarged stigma from postmarginal vein about 17 
times its height. 

Gaster slightly longer and distinctly broader than thorax, itself about 1 -6 times as long as 
broad. First tergite nearly as long as three following tergites combined. Pubescence of 
gaster mainly dark, relatively short, also bristles of cerci subequal in length. Epipygium in 
median line hardly longer than median length of sixth or fifth tergite. 

o*. Not known. 

Biology not known. Collected by sweeping in mixed forest. 
Holotype $. East Germany: Isle of Riigen, Baabe, vii.1960 (Boucek); presented 
to BMNH. 

Paratype. 1 $, collected with the holotype; in NM Prague. 

Named after Riigen = Ruj ana, old Slavonic name of the Isle. 

This is the most distinctive species of the genus, differing from all other European 
species mainly by the large body with extremely short pubescence on head and 
thorax, the antenna with large area of micropilosity in female, rather short gaster 
and not having one cereal bristle unusually long. Graham (1969 : 254, 255) 
mentions this species as 'sp. indet.' and states that the area of micropilosity is on 
ventral face of the clava (as it is commonly in Pteromalidae), while it is in fact placed 
dorso-apically. 



292 Z. BOUCEK 

PTEROMALINAE 
VELTRUSIA gen. n. 

Type-species: Veltrusia rara sp. n. 

Occiput not margined, temples and genae terete. Eyes with extremely fine inconspicuous 
hairs. Scrobes fairly deep but not sharply delimited though slightly angulate in front of 
ocellus; interantennal callus distinctly raised and extending into scrobes and towards clypeus. 
Lower face not protuberant. Clypeus mainly minutely reticulate, smooth at lower margin, 
subhexagonal : upper margin finely groove-like, short, horizontal, then obliquely descending to 
moderately deep tentorial pits, then converging downwards as broad shallow grooves; lower 
margin arched, slightly produced. Mouth margin sublaterally simple, arched; malar space 
convex. Left mandible 3-toothed. Lower edges of antennal toruli in lower ocular line. 
Antenna in female 13-segmented, with two anelli; pedicellus hardly longer than first funicle 
segment; funicle filiform, with segments decreasing in length; clava slender, bluntly pointed, 
both sutures perpendicular. Flagellum with pubescence short, inconspicuous, each segment of 
funicle and clava with one row of sensilla. 

Thorax elongate, not depressed, dorsally as well as head mainly rugose-reticulate, clothed 
with distinct and fairly dense dark hairs. Pronotum narrower than mesoscutum, collar sharply 
margined, hind margin emarginate with broad smooth belt; lateral panel shallowly concave, 
relatively broad. Notaular furrows complete, moderately deep. Scutellum broadly meeting 
mesoscutum, axillar furrows intercepted well inside of notauli; frenal furrow rather deep but 
irregular, wavy; frenum taking up apical quarter, its sculpture coarse longitudinal, raised 
reticulation; furrows of axillulae diverging, anteriorly deep, weak posteriorly. Dorsellum 
forming transverse crest behind basal crenulae, its posterior face reticulate. Propodeum 
reticulate, with coarse rugae in deep places; median carina rather irregular though high and 
raised anteriorly into a triangle which is blunt at top; plicae sharp, arched, slightly converging 
and high posteriorly; nuchal strip crescentic, irregular but distinct, separated from disc by deep 
cross-furrow with coarse crenulae; hind corner formed by sharp vertical irregular edge; pubes- 
cence of callus moderately long, not dense, whitish; supracoxal flange not conspicuous. Meta- 
pleura, mesepisternum and lower epimeron retiulate, upper epimeron smooth and reaching far 
down along the vertical subdividing furrow which is more fovea-like above where it turns 
towards metapleura. Presternum with mesolcus. Hind coxa reticulate, dorsally bare. Hind 
tibia with outer spur only half as long as the inner one. Legs relatively slender, basitarsi 
generally as long as two following segments combined. Fore wing pubescence dense on disc, 
marginal fringe short; basal fold hairy, cubital fold hairy except at proximal half of basal cell, 
the latter also with a few hairs below submarginal vein and on disc ; speculum broad but closed 
below (Text-fig. 34). Veins slender, marginal vein subequal to the postmarginal and nearly 
twice as long as the straight stigmal vein; the latter hardly knobbed. Parastigma rather stout, 
ending in a pale break. 

Gaster of female long, convex, lanceolate, posteriorly and laterally with dark hairs. Petiole 
very short, hidden, nearly smooth. First tergite moderately long, its hind margin sub- 
angularly produced in median two-quarters; tergites 2 to 4 subequal (Text-fig. 33), with hind 
margins nearly straight, the fifth tergite slightly longer than the fourth, the sixth still longer; 
one bristle of cercus slightly but not unusually longer. Ovipositor sheaths very slightly pro- 
truding. Hypopygium not reaching middle of gaster. 

o*. Unknown. 

Named after Veltrusy, a little town with old parkland in Bohemia, north of 
Prague, where the type-species and the two following new species (of Strejcekia 
gen. n.) were collected. 

This is another aberrant member of Pteromalinae and as such is not easy to place 



EUROPEAN PTEROMALIDAE 



293 



in the existing keys, for, because of complete notauli, it runs in them (Boucek in 
Peck, Boucek & Hotter, 1964; Graham, 1969) to Miscogasterinae. From the latter 
subfamily it differs in having the postmarginal vein as long as the marginal, and 
from most other genera by the pronotum which is rather short and carinate in the 
middle. 




Figs 33-37. 33-34- Veltrusia vara. 33, $ body; 34, part of fore wing. 35-36, Strejcekia 
elegans. 35, body of $ with fore wing; 36, head of ? in facial view. 37, Strejcekia 
brevior, body of $. (Hind margin of fifth tergite omitted in Fig. 35). 



294 Z. BOUCEK 

From most genera of Pteromalinae Veltrusia gen. n. differs in having the notauli 
clearly complete. In Graham's key it runs near to Dorcatomophaga Kryger, 
together with Strejcekia gen. n. described below. Both these new genera differ from 
Dorcatomophaga by several characters which may be summed up in the following 
way, altering the key by Graham on p. 360, couplet 41 (40). 

41 (40) Notauli complete, distinctly impressed throughout except sometimes just at 
the hind margin of mesoscutum; hind corners of propodeum with vertical 
ridge jutting over the base of hind coxa, the corner in dorsal view usually 
sharp, rectangular or acute; gaster usually convex . . . . .41a 

- Notauli almost always incomplete and reaching at most somewhat more than 

half way across the mesoscutum; very rarely traceable to its hind margin 
but then very superficial posteriorly, and propodeum not sharp-angled 
when viewed from above ......... 42 

41a (41) Pronotal collar with sharp carina; centres of antennal toruli above level of 
ventral edge of eyes; postmarginal vein about as long as the marginal; 
plicae of propodeum sharp and high, though slightly irregular; median carina 
raised to a tooth anteriorly ...... VELTRUSIA gen. n. 

Collar rounded, without distinct carina; antennal toruli below the lower 

ocular line ; postmarginal vein shorter than the marginal . . . .41b 

41b (41a) Antennal scrobes shallow, without subdividing median crest; lower face 
radiately striate and bearing only short, inconspicuous hairs; eye longer 
than malar space; pronotum in dorsal view not forming angular shoulders; 
marginal vein slightly thickened; propodeum shallowly reticulate-punctured, 
median carina and plicae obliterated ; gaster alutaceous all over. 

DORCATOMOPHAGA Kryger 

- Scrobes deep and in lower half (or more) separated by the median crest ; lower 

face not striate but clothed with conspicuous long hairs; pronotum in dorsal 
view with subrectangular shoulders, though much narrower than meso- 
scutum; marginal vein slender ; propodeum very deeply reticulate-punctured , 
dull (Text-fig. 35), with high crest-like plicae and median carina, the latter 
raised and widened anteriorly; gaster at least anteriorly mainly smooth. 

STREJCEKIA gen. n. 

Veltrusia rara sp. n. 

(Text-figs 33, 34) 

?. Black with dark green or dull bronze tinge, which is slightly more vivid on vertex and 
thoracic dorsum. Scapes and legs beyond coxae mainly brownish testaceous, femora more or 
less infuscate, trochantins and knees paler. Mandibles and palpi fuscous. Pedicellus fuscous, 
flagellum black. Wings subhyaline, veins brown. Length 4 mm. 

Head fully i-i times as broad as mesoscutum, fully twice as broad as long (52 : 25) in dorsal 
view, with temples moderately receding, only one-fifth the length of eyes. POL to OOL as 
11 -5 to 6-5, lateral ocellus slightly nearer to the anterior one than to occiput; relative width of 
vertex 30, eye 25 : 19-5, malar space 14, mouth width 21, height of head 43, width 52. Inner 
eye orbits very slightly diverging downward. Scrobes above ending about one diameter from 
ocellus. Fine reticulation behind malar sulcus engraved, alutaceous. Relative length of 
scapus 22, flagellum plus pedicellus 60, i.e., 1-15 times width of head. Pedicellus dorsally 
twice as long as broad, distinctly narrower than f unicle ; both anelli combined about as long as 
broad; first funicle segment 1-5 times, the sixth 0-9 times as long as broad, clava 2-2 times as 
long as broad. 



EUROPEAN PTEROMALIDAE 295 

Thorax (Text-fig. 33) from collar margin down to apex of propodeum about 1 -6 times as 
long as broad. Relative length of collar in the middle 4, at sides in dorsal view 12, length of 
mesoscutum in middle 30, scutellum 26. Mesoscutum finely transversely reticulate, with bases 
of hairs raised. Scutellum on disc minutely engraved-reticulate, with scattered piliferous 
punctures, part between axillulae and in front of frenum as long as broad, axillulae moderately 
sloping, rather broad. Propodeum medially half as long as scutellum, the raised nuchal strip 
confined to apical quarter. Oval spiracles removed by their longer diameter from metanotal 
margin. Fore wing relative length 154, width 61, costal cell 52 (width about 5), marginal 
vein 36, postmarginal 36, stigmal 20, distance between stigma and postmarginal vein 8. Lower 
surface of costal cell hairy, hairs proximally reduced to incomplete double line. 

Gaster nearly 1-3 times as long as head plus thorax combined, 3-1 times as long as broad 
itself, broadest in basal third. First tergite slightly shorter than three following tergites com- 
bined, the fourth with hind margin slightly emarginate; first tergite smooth, laterally near 
base with a few hairs, the following tergites basally with obliterated alutaceous sculpture, the 
fifth and sixth slightly more distinctly engraved-reticulate with raised piliferous punctures 
(except at hind margin). 

0*. Unknown. 

Biology unknown. Judging from the taxonomic characters the species may be 
a parasite of xylophagous beetles, most probably of Anobiidae. The specimen, 
along with the two species of Strejcekia described below, was beaten from bushes and 
trees in an old park. 

Holotype $. Czechoslovakia: Bohemia, Veltrusy, 9. v. 1959 (/. Strejcek); in 
Boucek Collection. 

Together with Dr Strejcek and my assistant we have been trying to get some more 
material of these interesting forms, which seem to be very rare (hence the specific 
name), but without success. In the meantime I consulted various colleagues and 
we agreed that the three specimens, collected in one spot, on the same day, belong 
to two new genera and to three new species. It is only now, after more than ten 
years, that I am publishing their descriptions. 

STREJCEKIA gen. n. 

(Text-figs 35-37) 
Type-species: Strejcekia elegans sp. n. 

Body hardly metallic; head and thorax reticulate, reticulation mainly obliterated dorsally 
but very deep on propodeum; gaster nearly smooth. Head with eyes relatively small, their 
pubescence short, not conspicuous. Occiput not margined but with unusually coarse rugose 
reticulation; temples and genae terete. Scrobes not margined, rather deep; interantennal 
crest high, reaching narrowly far into scrobes and also downward, as convex supraclypeal area. 
Lower face not protuberant, distinctly hairly, not radiately striate. Clypeus very narrow, 
ill-defined above, its lower margin more or less produced and thin, surface not striate. Sub- 
lateral margins of mouth not sinuate. Mandibles small, normal, 3-toothed, upper tooth tran- 
cate. Antennae in female not very long (Text-figs 35, 37), 13-segmented, inserted below lower 
ocular line, not far below middle of face. Scapus slender, distinctly longer than eye ; pedicellus 
longer than first funicle segment; two short anelli: nagellum weakly clavate, the six segments 
of funicle gradually decreasing in length, each segment narrowing basad and generally with 
one row of sensilla; three-segmented clava blunt at apex, with sutures almost perpendicular. 

Thorax not depressed, elongate. Pronotum much narrower than mesoscutum, collar only 
bluntly set off (not carinaceous) , in dorsal view with distinct angular shoulders, in middle short 



296 Z. BOUCEK 

but side panels rather long, shallowly concave. Notauli complete, not very shallow. Scuto- 
scutellar suture nearly straight, separated from scutellum by a cross-furrow. Scutellum 
reticulate, frenal furrow marked by deep bases of irregular longitudinal grooves of frenum. 
Dorsellum short, very deeply reticulate-punctured as is the propodeum, dull. Propodeum 
with broad, ridge-like median carina rising into a blunt tooth anteriorly and with more or less 
distinct, equally ridge-like, subparallel plicae; spiracles small, oval; part laterad and caudad 
of spiracle in addition to the reticulation coarsely and irregularly rugose; callus with irregular 
longitudinal ridge; hind corners of propodeum jutting over base of metacoxa and with a vertical 
ridge, but not nearly reaching level with the protruding median part of propodeum, which 
forms, however, no neck; no conspicuous supracoxal flange. Prepectus large, with distinct 
vertical ridge anteriorly, depressed on disc. Mesopleura and metapleura mainly deeply 
reticulate; mesepimeron with deep fovea above the middle, in front of fovea with a vertical 
strip of shallower sculpture. Hind coxa reticulate, dorsally bare. Legs fairly slender, hind 
tibia with only one distinct spur. Basitarsi of all legs long, dorsally fully as long as following 
two segments together. Fore wing moderately densely hairy, marginal fringe of medium length ; 
costal cell unusually narrow; basal fold hairy but speculum open or closed below; veins slender, 
marginal vein at most as long as the postmarginal, much longer than the stigmal which is 
hardly knobbed. 

Gaster in female (Text-figs 35, 37) convex, conically lanceolate, nearly smooth and only 
poorly pubescent posteriorly. Petiole short and mostly hidden under the propodeum, but with 
several coarse longitudinal rugae. Hind margins of tergites mainly straight, the first and 
second relatively large though not reaching middle of gaster. Epipygium not very long, with 
a group of denser short hairs in front of the cercus, which has one bristle conspicuously longer 
than the others. Ovipositor sheaths hardly protruding. Hypopygium reaching middle of 
gaster. 

Males not known. 

Named in honour of Dr J. Strejcek of Prague, a keen entomologist working in 
nature conservation, to whom I am indebted for some very interesting material of 
Chalcids. 

Strejcekia gen. n. also comes near to Dorcatomophaga Kryger and its distinguishing 
characters are summed up above along with Veltrusia gen. n. 

Key to Species 

Females 

1 Body slender (Text-fig. 35), thorax dorsally fully 2-3 times as long as breadth of 
pronotum ; antennae longer, flagellum plus pedicellus combined about 1 -4 times as 
long as breadth of head, funicle segments 1-4 not transverse; scrobes only moder- 
ately deep, not sharply delimited above and not reaching near to the median ocellus; 
interscrobal callus blunt, not very high (Text-fig. 36) ; median part of propodeum 
strongly protruding beyond posterolateral corners, the strong ridgelike plicae 
slightly longer than distance between them anteriorly . . elegans sp. n. (p. 297) 

- Body broader (Text-fig. 37), thorax dorsally only 1-9 times as long as breadth of 
pronotum ; antennae shorter, stouter, flagellum plus pedicellus combined distinctly 
shorter than breadth of head; funicle segments 1-6 strongly increasingly trans- 
verse; scrobes very deep, abruptly ending about one diameter before ocellus, 
interscrobal crest narrow, high and sharp ; median part of propodeum only slightly 
protruding beyond level of posterolateral corners (above base of coxa) ; weak 
plicae only about two-thirds as long as distance between them anteriorly. 

brevior sp. n. (p. 297) 



EUROPEAN PTEROMALIDAE 297 

Strejcekia elegans sp. n. 

(Text-figs 35, 36) 

$. Body black, dorsally with very slight dark green tinge. Legs, base of antennae and 
tegulae dark testaceous; fiagellum except anelli fuscous, also coxae infuscate basally. Wings 
subhyaline, venation pale testaceous. Length 2-4 mm. 

Head in dorsal view broader than mesoscutum as 35 : 30, than the pronoum as 35 : 22-5, 
itself 1-75 times as broad as long, with temples about three-quarters the length of eyes but 
rather strongly arcuately receding; occiput deeply emarginate, taking up slightly more than 
half the head breadth. POL equals OOL. Frons laterally strongly convex. Scrobes moder- 
ately deep, above ending nearly two diameters from the ocellus, but not well delimited. Inter- 
antennal callus (Text-fig. 36) blunt and fading out slightly above middle of scrobes. Eyes 
prominent though small, with distinct short pubescence. Relative measurements: head width 
35, height 31, width of frons 26, oval eye 12 : 9, malar space 13-5, mouth width 20, distance 
between lower margin of clypeus and antennal toruli 11-5, scapus length 17, fiagellum plus 
pedicellus 49, i.e., 1-4 times the width of head. Scapus reaching level with lower edge of 
median ocellus; pedicellus dorsally 2-4 times as long as broad and as long as anelli plus first 
funicle segment; both anelli together shorter than broad; first funicle segment with sensilla 
confined to distal half, basally constricted, about 1-5 times as long as broad; the fifth and sixth 
segment slightly transverse. Clava about 2-2 times as long as broad, apical nipple with small 
area of micropilosity. 

Thorax measured from anterior margin of collar to apex of propodeum nearly 1 -8 times as 
long as breadth of mesoscutum, fairly convex. Hind margin of pronotum smooth, deeply 
emarginate. Mesoscutum and scutellum distinctly hairly, fairly shiny as the reticulation is 
rather fine and shallow. Mid lobe of mesoscutum strongly convex, 11 times as broad as long. 
Scutellum slightly elongate, at apex subtruncate; frenum taking up more than apical one-quarter, 
coarsely longitudinally rugose; disc of scutellum nearly smooth, very finely alutaceous, with 
scattered fine piliferous punctures. Axillulae short, moderately sloping, dull, deeply reticulate. 
Dorsellum of metanotum dull, deeply reticulate-punctured as is the propodeum. Apex of 
protruding median part of propodeum emarginate, not margined; median ridge triangularly 
expanding and rising towards base; spiracle very small, round; lateral callus with a longitudinal 
crest, the hairs thin and not dense. Hind femur nearly 6 times as long as broad, clothed 
sparsely with thin and long hairs. Wings hardly exceeding apex of gaster. Fore wing narrow 
(Text-fig. 35), regularly rounded at apex, relative length no, width 40, costal cell 43 : 3, 
marginal vein 25, postmarginal 20, stigmal vein 10. Lower surface of costal cell with only one 
row of hairs, this precurrent; submarginal vein smoothly joining parastigma; anterior margin 
of wing with marginal and postmarginal vein forming a smooth arch; stigmal vein angle about 
45 , stigma very small, subtriangular, the short uncus almost parallel to postmarginal vein. 
Base of wing almost all hairy but hairs sparse, rather long; speculum very small. Hind wing 
relative length 85, width 19, longest fringe 3; fairly broadly rounded at apex. 

Gaster narrower than mesoscutum, about 1 -2 times as long as head plus thorax combined, 
itself 3-2 times as long as broad, smooth, posteriorly bearing some sparse long thin hairs. 

o*. Not known. 

Biology not known. 

Holotype $. Czechoslovakia: Bohemia, Veltrusy, 9.V.1959 (Strejcek); in 
Boucek Collection. 

Strejcekia brevior sp. n. 

(Text-fig. 37) 
$. Black; propodeum and base and apex of gaster slightly brownish; scapes and pedicels 



298 Z. BOUCEK 

and legs apart from coxae mainly pale testaceous, pedicels and femora slightly infuscate. 
Wings subhyaline, venation light brown. Length 2-7 mm. 

Head slightly broader than mesoscutum (as 42 : 38; Text-fig. 37), in relation to pronotum 
as 42 : 31, itself in dorsal view 1-75 times as broad as long, with temples about 0-7 the length 
of eyes. POL to OOL as 11 : 8. Frons on sides strongly protuberant, rounded; scrobes deep, 
narrowing above and there in the middle angulately delimited, only about one diameter in 
front of ocellus; interscrobal crest narrow, high, sharp up to half of scrobes, more dorsally 
much lower but still distinct. Lower face below antennal toruli subhorizontally rugose-striate, 
irregularly rugose nearer to mouth. Also gena dull, deeply irregularly rugulose, malar sulcus 
below replaced by a blunt ridge. Relative measurements: width of head 42, height 36, width 
of frons 30, oval eye 15 : 11, malar space 17, mouth width 18, scapus length 18, flagellum plus 
pedicellus 37, i.e., o-88 the head width. Pedicellus dorsally hardly 1-5 times as long as broad; 
first funicle segment about 1-5 times, the sixth about twice as broad as long; clava less than 
twice as long as broad. 

Thorax dorsally from anterior corners down to apex of propodeum 1-57 times as long as 
breadth of mesoscutum. Sculpturally similar to S. elegans but all parts shorter, broader (hence 
the specific name) ; scutellar frenum posteriorly reticulate; mid lobe of mesoscutum only weakly 
convex, anteriorly cross-striate-alutaceous; notauli shallower but clear-cut down to scuto- 
scutellar suture. Scutellum as long as its maximum breadth measured between axillulae. 
Dorsellum truncate when seen from in front or from behind, with sublateral parts rather high. 
Propodeum in middle two-thirds the length of scutellum, median carina in anterior two-thirds 
replaced by large triangular blunt tooth the apex of which is in line connecting the postero- 
lateral corners of propodeum; these blunt in dorsal view but formed by short, vertical ridge. 
Callus inconspicuously hairy. Hind femur 5 times as long as broad, dorsally with short 
pubescence, only on lower edge at apex withsome longishhairs. Forewing relative measurements: 
length 112, width 41, costal cell 44, marginal vein 24, postmarginal 24, stigmal vein 11. Basal 
cell bare; speculum of medium size, reaching broadly cubital hair-line; posterior corner of fore 
wing blunt but distinct, at level with stigmal vein, apex of wing thus asymmetrically rounded, 
more strongly so anteriorly than posteriorly (unlike in 5. elegans). 

Gaster as broad as mesoscutum, 1-12 times as long as head plus thorax combined, itself 
about 2-2 times as long as broad. First tergite the longest, anteriorly on sides with a loose 
patch of longish hairs, apex of gaster with relatively short hairs, epipygium with very short 
hairs; second tergite shorter than the first as 16 : 22, but nearly as long as three following 
tergites (3-5) together. 

<J. Not known. 

Biology not known, but as in the preceding species, the morphological affinity 
suggests parasites of xylophagous beetles, probably with some rather cryptic way of 
life, as may be judged from the relatively small eyes and, at least in Strejcekia 
elegans sp. n., from the rather long and thin hairs on the gaster. 

Holotype $. Czechoslovakia: Bohemia, Veltrusy, 9. v. 1959 (Strejcek); in 
Boucek Collection. 

RHIZOMALUS gen. n. 

(Text-figs 41, 42) 
Type-species: Rhizomalus cwpreus sp. n. 

Head and thorax very finely and densely, but shallowly, reticulate, very shortly and fairly 
densely hairy, piliferous punctures very distinct. Head in dorsal view moderately transverse; 
occiput slightly emarginate, not margined; frons convex, scrobes distinct but not very deep 



EUROPEAN PTEROMALIDAE 299 

Lower face not protuberant, finely radiately reticulate to striate (Text-fig. 41); genae convex, 
posteriorly rounded. Clypeus small, tentorial pits and upper margin not distinct, the lower 
margin truncate. Left mandible 3, the right 4 teeth, not large, moderately curved. Antennae 
in both sexes rather short, 13-segmented, inserted slightly above lower ocular line but below 
centre of face. Scapus hardly broadening distally, sublinear, not nearly reaching median 
ocellus; pedicellus longer than first funicle segment; two short anelli; flagellum rather stout, 
sublinear, all funicle segments transverse, almost equal in length, each with one row of longi- 
tundinal sensilla, in $ very shortly haired, hairs longer in £; clava ovate, bluntly pointed, its 
sutures perpendicular, micropilosity area reduced to terminal nipple. 

Pronotum rather short, collar in middle sharply carinaceous; in dorsal view not forming 
shoulders on sides. Mesoscutum with notauli indicated as superficial lines on anterior half; 
scuto-scutellar suture weakly sinuate laterally. Scutellum slightly convex, frenal groove 
barely perceptible. Metanotum linear. Propodeum very short, finely rugulose-reticulate, 
transversely convex; nucha represented by a narrow elevated strip; median carina vague or 
weak, plicae absent; callus sparsely hairy; spiracles small, short-oval, removed from meta- 
notum by more than their diameter. Prepectus small, triangular, weakly reticulate, anteriorly 
without carina. Mesepimeron with distinct arched subdividing furrow, upper epimeron 
smooth, the lower more strongly reticulate than outer surface of hind coxa; the latter dorsally 
bare. Legs rather stout, hind femur only slightly more than 3 times as long as broad; hind 
tibia longer than femur, with one spur; mid tibia with spur longer than width of tibia at apex, 
slightly shorter than basitarsus dorsally. Fore wing pilosity dense and very short (Text-fig. 40), 
marginal ciliation developed throughout; costal cell broad; parastigma slightly thickened and 
distally with a pale break; marginal vein very slightly thickened, hardly shorter and much 
broader than the postmarginal, slightly longer than the stigmal vein; the latter with small 
stigma. Basal cell mainly open below but more or less hairy at the basal hair-line; speculum 
not reaching marginal vein, mostly open below. 

Gaster sublanceolate, dorsally mostly depressed. Petiole very short, hidden, smooth. 
Basal tergite with hind margin mainly entire. Bristles of cerci subequal in length to the 
normal pilosity. Ovipositor not exserted. Tip of hypopygium situated about half way along 
the gaster. 

In male, mouth with labiomaxillary complex and legs normal, as in female. 

The genus Rhizomalus seems to be nearest to Hobbya Delucchi and Cecidostiba 
Thomson, both morphologically and biologically. In Graham's key to the genera 
of the European Pteromalinae (1969 : 353-409) the female runs to couplet 132 and 
may be keyed out there in the following way. 

132 (131) Fore wing broadly suffusedly infumate below marginal vein, the latter 
slightly thickened, shorter than the postmarginal and little longer than 
the stigmal vein; stigma very small; basal cell hairy in distal quarter or 
more. Propodeum short, plicae indistinct. Antennae very short, all 
funicle segments transverse; pedicelus about twice as long as the first 
funicle segment RHIZOMALUS gen. n. 

Fore wing hyaline or with different fuscous markings; marginal vein not 
thickened and not shorter than the postmarginal vein; basal cell usually 
bare; stigma either relatively large or propodeum about half the length 
of the scutellum. Antennae much longer, basal segments at least slighty 
elongate ............ 132a 

132a (132) Here Graham's couplet 132, keying out Cecidostiba Thomson and 
Nephelomalus Graham 



300 Z. BOUCEK 

The male runs in Graham's key to couplet 128 (p. 404), partly to Pegopus Forster, 
but differs from that genus as follows. 

a Antennae short, scapus not nearly reaching the ocellus, all funicle segments distinctly 

transverse; left mandible 3-toothed .... RHIZOMALUS gen. n. 

- Antennae not so short, scapus reaching to the level of vertex, funicle segments sub- 
quadrate or slightly elongate; both mandibles 4-toothed . . PEGOPUS Forster 

Rhizomalus cupreus sp. n. 

(Text-figs 41, 42) 

$. Cupreous; head and thoracic dorsum more reddish, gaster basally more brownish metallic, 
apically purplish. Antennal base including anelli, then trochanters, knees, tibiae and tarsi 
testaceous; antennal flagellum blackish brown; coxae concolorous with thorax, femora 
extensively fuscous with metallic tinge, sometimes also tibiae slightly infuscate or gaster 
beneath brownish. Fore wing, except for subhyaline basal third, more or less yellowish to 
brownish infumate, more distinctly so on disc below marginal and stigmal vein; venation 
brown. Hairs on thoracic dorsum mainly dark. Length 2-4-3 mm. 

Head distinctly broader than mesoscutum (51 : 44), in dorsal view 1 9 times as broad as long, 
i.e., rather stout, with temples moderately receding and hardly one-third as long as eye. POL 
1-35 times OOL; ocellar triangle 2-2 times as broad as high. Eyes not prominent, with ex- 
tremely short sparse hairs. Supraclypeal area slightly convex but not well delimited. Clypeus 
flat, depressed, more strongly receding towards mouth than the adjoining face, its lower margin 
smooth. Mouth margin thin, regularly arched. Relative measurements: head width 51, 
height 42, width of frons 35, eye 23 : 17-5, malar space 14, width of mouth 25, distance be- 
tween upper edge of antennal toruli and median ocellus 20, flagellum plus pedicellus combined 
39, i.e., 0-77 the head width. Pedicellus dorsally about 1-7 times as long as broad; flagellum 
hardly clavate (Text-fig. 41); first funicle segment 1-05-1-2 times as broad as long, the sixth 
about i-6 times as broad as long; clava as long as 2-5 preceding segments combined. 

Pronotum with collar in middle about 1/8 the length of mesoscutum, its carinaceous anterior 
margin weakly arched, sides strongly diverging, slightly bulging posteriorly. Scutellum 
i-i times as long as maximum breadth less axillulae; frenum relatively shiny, its reticulation 
wide-meshed, obliterated; scutellar disc extremely densely reticulate, front margin meeting 
mesoscutum for one-quarter of mesoscutal breadth, as broadly as each axilla. Propodeum 
duller than disc of scutellum, in the middle less than one-third the length of scutellum; postero- 
lateral corners rounded, with a small supracoxal flange. Fore wing broad (136 : 53), marginal 
vein 8-2-9-5 times as long as broad. Relative length of costal cell 46, marginal vein 22, post- 
marginal 24, stigmal vein 18 (Text-fig. 40). 

Gaster 1 -05-1 -25 times as long as head plus thorax combined. First and second tergite in 
middle of hind margin sometimes submarginate, dorsally nearly smooth, the following tergites 
alutaceous basally. Weak pubescence on sides and apex of gaster dark. 

o*. The two specimens available are unusually small, i-6 mm. Head perhaps therefore 
relatively broader, 1-2 times as broad as mesoscutum. In colour very similar to female, but 
infumation of the fore wing weak. Flagellum plus pedicellus combined o-86 the breadth of 
head; pedicellus dorsally scarcely 1-5 times as long as broad; all funicle segments distinctly 
transverse, clothed with semidistant hairs which are nearly as long as segments. Gaster hardly 
longer and much narrower than the thorax. 

Biology. Reared from oak gall of the Cynipid Andricus quercusradicis (¥.). 

Holotype $. France : C6tes-du-Nord, Erquy-les-Bains (H. B. Preston) ; in 
BMNH. 



EUROPEAN PTEROMALIDAE 



301 




Figs 38-45. 38-40. Peridesmia montana. 38, body of $ with sculpture of propodeum 
indicated; 39, head of o"; 40, part of fore wing of $. 41-42. Rhizomalus cupreus. 41, 
head of $ in facial view; 42, fore wing venation in $. 43—45. Pteromalus paludicola. 
43, part of fore wing with venation and pilosity; 44, mouth region with mandibles; 
45, $ flagellum with pedicellus. 



302 Z. BOUCEK 

Paratypes. Britain: England, Oxfordshire, Bald Hill near Lewknor, i $, 
15.vii.1960 (Graham); Graham Collection. Czechoslovakia: Slovakia, Kovacov 
near Sturovo, 1 $, 17.vii.1969 (Boucek). Hungary: Baja, ex gall of Andricus 
quercusradicis, 1 $, 6.viii.i96i (Fekete). Yugoslavia: Dalmatia, Biograd na mora, 
1 $, vii.1968 (Boucek). Bulgaria: Sandanski district, 3 £, vi.1969 (Kocourek). 
Greece: Kalamaria near Thessaloniki, 4 $, 2 <J (one allotype), 1917 (/. Waterston), 
BMNH. Also 2 $ from the Moldavian S.S.R. (U.S.S.R.) examined but data not 
noted. Partly in Boucek Collection, partly in NM, Prague and BMNH, London. 

PTEROMALUS Swederus 

Pteromalus Swederus, 1795 : 201. Type-species: Ichneumon puparum Linnaeus; designated by 
Westwood, 1839. 

Pteromalus paludicola sp. n. 

(Text-figs 43-45) 

$. Bluish green; antennae blackish brown, scapes testaceous, infuscate at apex; legs except 
metallic coxae mainly reddish testaceous, but femora mostly infuscate, as well as fore tarsi and 
apex of mid and hind tarsi, sometimes also mid and hind tibiae infuscate. Wings hyaline, 
venation brown. Length 1-8-2-2 mm. 

Head about 1-28 times as broad as mesoscutum, dorsally about 2-15 times as broad as long, 
temples about half the length of the eyes, converging moderately and rather straight. POL 
about 1 1 times the OOL, the latter equals about 3 diameters of lateral ocellus. Head in facial 
view transversely subelliptic about 1-23 times as broad as high; lower face with clypeus dis- 
tinctly radiately striate (Text-fig. 44), lower margin of clypeus shallowly emarginate, medially 
depressed. Mandibles clearly 4-toothed. Genae slightly swollen just at mouth margin which 
is depressed from below at mouth corners and bordered by a groove sublaterally (seen from 
below). Relative measurements; width of head 50, of frons 32, eye 22 : 16, malar space 12, 
width of mouth 21, distance between lower margin of clypeus and antennal toruli 15, scapus 
17-5, flagellum plus pedicellus 38-5, i.e., about 0-75-0-8 the breadth of head. Scapus relatively 
short, not nearly reaching to ocellus; pedicellus dorsally about 1-5 times as long as broad, 
distinctly longer than first funicle segment (Text-fig. 45) ; the latter slightly transverse, the 
following segments more transverse, sixth funicle segment about 1-4 times as broad as long; 
clava ovate-subacuminate, about as long (0-95-1-04) as three preceding segments combined. 
Flagellum hardly to slightly clavate, each funicle segment with one row of sensilla. 

Thorax about 1 -5 times as long as broad, dull, densely punctured-reticulate, rather unusually 
densely clothed with dark short hairs (conspicuous in lateral view). Dorsum not flattened. 
Pronotum 0-82 times as broad and medially one-sixth to one-seventh as long, as mesoscutum. 
The latter 1 -74 times as broad as long, on disc with reticulations as dense as on scutellum but 
elsewhere more finely reticulate. Scutellum rather flat, hardly as long as broad, frenum not 
distinctly marked off sculpturally. Propodeum half as long as scutellum; median carina 
indicated only basally; plicae posteriorly at base of nucha very low; hairs of callus partly dark, 
rather dense also behind spiracle; depressed part above hind coxa and behind callus rather 
short. Legs moderately stout, hind femur 4-5 times as long as broad, hind tibia about 1-2 
times as long as hind tarsus. Fore wing (Text-fig. 40) unusually extensively hairy, basal cell 
hairy all over, speculum usually closed below or nearly, lower surface of costal cell even basally 
with double or triple row of hairs. Relative lengths of veins: marginal 21, postmarginal 25, 
stigmal 15; in a paratype m : pm : st as 18 : 25 : 14. 

Gaster about as long or slightly shorter than thorax, about 1 5-1 -6 times as long as broad, 
dorsally depressed. Basal tergite occupying slightly less than one-third the total length. 

o\ Not known. 



EUROPEAN PTEROMALIDAE 303 

Biology. Host not known. All specimens were collected in marshy habitats 
(hence also the specific name) . 

Holotype $. Czechoslovakia: Bohemia, ftevnicov, 14.viii.1958 {Boucek); 
presented to BMNH. 

Paratypes. Czechoslovakia: Bohemia, Bfehyne near Doksy, 5 $, 17.vii.1963 
(Boucek) ; partly in Boucek Collection, partly in NM, Prague. 

Pteromalus paludicola sp. n. with its completely hairy basal cell can be easily 
separated from all the other European species of Pteromalus and Habrocytus, most 
of which are keyed in a combined key by Graham (1969 : 495-523). This character 
occurs otherwise only in some Pteromalus venustus Walker, as discussed by Boucek 
(1970 : 74) and, possibly, in the rather enigmatic P. vopiscus Walker (see Graham, 
1969 : 492), which differs in having fairly convex scutellum. Otherwise both can 
be separated from P. paludicola as follows. 

1 Length of flagellum plus pedicellus only o-8 the width of head, all funicle segments 
transverse; clava about as long (0-95-1-04) as three preceding segments combined. 
Basal cell completely hairy, speculum mostly closed below paludicola sp. n. (p. 302) 

- Flagellum plus pedicellus as long as 0-86-0-93 the width of head, proximal funicle 
segments not transverse; clava 1-24-1-3 the length of two preceding segments 
combined. Basal cell proximally bare, speculum open below. 

venustus Walker and vopiscus Walker 

Another species similar to Pteromalus paludicola sp. n. is Habrocytus crassicornis 
(Zetterstedt) occurring in Czechoslovakia in the same habitats. It has similarly 
hairy wings and short antennae, but the left mandible is 3-toothed, lower face 
strongly striate, scutellum strongly convex, venation different, etc. 

PERIDESMIA Forster 

Peridesmia Forster, 1856 : 65. Type-species: Isocyrtus (Trichomalus) aquisgvanensis Mayr; 
designated by Gahan, 1923. 

Peridesmia montana sp. n. 

(Text-figs. 38-40) 

$. Dark green, in places with bronzy or cupreous tinge; antennal scapes and pedicels as 
well as legs beyond coxae, testaceous; flagellum blackish. Wings subhyaline, venation testa- 
ceous. Length 2-8-3 mm. 

Head fully 1 -4 times as broad as mesoscutum (Text-fig. 38), in dorsal view 2-1 times as broad 
as long, with temples slightly less than one-third the length of compound eyes. POL 0-9-0-95 
the OOL. Relative width of frons 37, width of head 64, eye 305 : 24, malar space 15-5, 
scapus 22. In facial view head 1-27-1-3 times as broad as high, with genae arched, strongly 
converging; mouth margin sublaterally strongly receding inward but distinctly produced at 
either side of clypeus. Flagellum plus pedicellus combined about o-86 the width of head; 
pedicellus dorsally fully twice as long as broad; flagellum clavate, second anellus only slightly 
transverse ; first funicle segment slightly broader than pedicellus, about 1 -5 times as long as 
broad, the sixth about 0-75 times the breadth and 1-5 times as broad as pedicellus; clava sub- 
acuminate, as long as 2 -5 preceding segments combined. 



304 Z. BOUCEK 

Thorax i-6 times as long as broad, rather finely reticulate-punctured. Pronotum distinctly 
narrower than mesoscutum, as 38 : 45 ; collar indistinctly edged, in dorsal view sides diverging, 
not protruding. Mesoscutum about 1 -8 times as broad as long, longitudinally moderately 
strongly convex. Scutellum fairly convex, as long as mesoscutum and slightly longer than its 
breadth measured posteriorly between axillulae. Propodeum medially 0-72 the length of 
scutellum ; median carina and plicae very strong and high, plicae strongly sinuate ; median area 
cordiform, fully 1 -6 times as broad as long, its bottom rather shiny, with some obliquely 
diverging rugae and only traces of reticulation, in deeper parts nearly smooth, posteriorly 
delimited by highly carinaceous angulate edge of nuchal strip (Text-fig. 38) ; the strip in form of 
a low arched trapezoid, depressed and scarcely strigulose on disc, its sides subparallel, raised, 
nearly as long as the strip in the middle. Lateral parts of propodeum beyond spiracles and 
beyond posterior half of plicae densely hairy. Fore wing densely hairy on disc but nearly bare 
in basal third, with only a few short hairs near upper part of basal fold (Text-fig. 40) ; lower 
surface of costal cell with complete hair-row, partly doubled basally, double or triple distally. 
Relative lengths of veins: marginal 25, postmarginal 23, stigmal 17. 

Gaster ovate-acuminate, depressed, slightly shorter than head plus thorax combined, broader 
than thorax but narrower than head. First tergite laterally densely hairy. 

<J. In colour similar to female but more vividly cupreous on vertex and thoracic dorsum; 
the smooth strip on head (Text-fig. 39) blackish purple. Antennal flagellum dark testaceous 
except for blackish distal third. Length 2-5 mm. 

Head very stout, 1 -6 times as broad as mesoscutum and dorsally 1 -9 times as broad as long, 
with the smooth strip very broad and long, touching the eyes and extending from the mouth 
corner up on temples and vertex forward to upper frons. POL about o-8 the OOL. Antennae 
only slightly more slender than in female, flagellum plus pedicellus combined o-8 the breadth 
of head, second anellus subquadrate, distal funicle segments subquadrate. Propodeum with 
median area still more shiny, with more distinct but sparser rugae, the reticulation traceable 
only on the disc; nuchal strip narrower than in female, its hind margin more raised. Gaster 
weakly convex, about three-fourths the length of thorax; first tergite covering one-half. 

Biology not known. All three specimens were collected by sweeping grass on 
montane meadows. 

Holotype $. Czechoslovakia: Slovakia, Remetske Hamre, 10.vii.1960 (Strejcek) ; 
in Boucek Collection. 

T 

Paratypes. Czechoslovakia: Bohemia, Krkonose Mts, Dolni Mala Upa, 1 $, 
8.ix.i968 (Boucek); Slovakia, Ulic-Stionka, 1 $ (allotype), 18.viii.1957 (L. Masner). 

Two European species were previously known and the new species may be 
separated from them mainly on the following characters. 

1 Median area of propodeum in both sexes rather shiny, with some diverging 

rugae, nearly smooth posteriorly in the depression in front of nuchal strip; 
the latter trapezoidal, its highly carinaceous sides hardly shorter than length 
of the strip in the middle, o*: smooth strip behind eyes very broad, reaching 
from mouth over temples and vertex forward on frons beyond level of front 
edge of median ocellus, dorsally strip as broad as POL. 

Pronotum distinctly narrower than the mesoscutum. Length of body 
2-3-3 mm ........ montana sp. n. (p. 303) 

- Median area of propodeum dull, punctured-reticulate; nucha either not set off 

posteriorly by a sharp cross-carina (P. congrua) or the nuchal strip is in form 
of a low triangle and pronotum is scarcely narrower than mesoscutum 
(P. discus). In o* the smooth postocular strip either much shorter or, if 
reaching vertex, much narrower than above . . . . . . ' 2 



EUROPEAN PTEROMALIDAE 305 

2 (1) For separation of P. congrita and P. discus see Graham (1969 : 701). 

Both Peridesmia congrua (Walker) and P. discus (Walker) are not uncommon in 
Czechoslovakia, but no new information on their biology is available. P. discus is 
known to me also from Yugoslavia: Kopaonik Mts, Milanov Vrh, 24.viii.1958 
(Jankovic) . 

SPANIOPUS Walker 
(Text-figs 46-55) 
Spaniopus Walker, 1833 : 466. Type-species: Spaniopus dissimilis Walker, by monotypy. 

For synonymy and references see Graham, 1969. 

Graham (1969 : 702-707) keyed out females of two species and males of four 
species. The mostly fresh material at my disposal contains seven European species, 
two of which have proved new to science. 

Key to European Species 
Females 

1 Eyes relatively small; in facial view width of frons about twice the height of 

eye, about equal to height of head (Text-fig. 58). Fore wing usually with 
several dispersed spots (Text-fig. 57) ; marginal vein about twice as long as 
the stigmal ........ polyspilus Graham (p. 307) 

Eyes larger; frons in facial view distinctly narrower than height of head. Fore 
wing markings otherwise or missing; marginal vein only rarely twice as long 
as the stigmal, mostly shorter ......... 2 

2 (1) Body relatively slender; thorax 1-6-1-67 times as long (from anterior edge of 

collar) as breadth of mesoscutum; gaster 1-72-2-1 times as long as broad, 
sublanceolate, sides of sixth tergite converging at an angle less than 50 . 
Notauli distinct as impressed lines along about 3/4 of mesoscutum. Fore 
wing usually with one lunate macula below (but not touching) stigmal vein 
(Text-fig. 62), sometimes with another faint subapical spot. Associated with 
Phragmites ........ peisonis (Erdos) (p. 307) 

Body stouter; thorax 1-45-1-57 times as long as broad; gaster shorter, ovate- 
acuminate, sides of sixth tergite converging at an angle of about 6o° or more. 
Notauli not reaching beyond o-6 along mesoscutum. Fore wing, if spots 
distinct, differently marked. Probably never associated with Phragmites . 3 

3 (2) Antennal pedicellus dorsally fully twice as long as broad, first funicle segment 

fully 1-5 times as long as broad and distinctly constricted basally; whole 
antenna fulvous. Marginal vein often more than 1 -75 times as long as the 
stigmal vein. Fore wing, if infumate, with a broad cloud below marginal and 
stigmal vein ............ 4 

- Pedicellus at most i-8 times as long as broad, first funicle segment at most 1-3 

times as long as broad, differing little in length from distal funicle segments; 
flagellum at least partly fuscous. Marginal vein at most 1 -75 times as long as 
the stigmal vein. Fore wing markings different (or missing) ... 5 



306 Z. BOUCEK 

4 (3) Fifth funicle segment subquadrate, the sixth subtransverse ; flagellum less 

strongly clavate, combined with pedicellus about as long as width of head. 
Fore wings larger, distinctly exceeding apex of gaster; marginal vein about 
1 -75 times as long as the stigmal, about 1 -5 times as long as the postmarginal 
which is strongly tapering apically; angle between inner margins of post- 
marginal and stigmal veins about 50 . . monospilus (Thomson) (p. 310) 

- Fifth and sixth funicle segments considerably transverse, flagellum more clavate 

(Text-fig. 49), its length combined with pedicellus only about 0-9 times the 
width of head. Fore wing smaller, about reaching apex of gaster; marginal 
vein about twice as long as the stigmal, about 1 -8 times as long as the post- 
marginal, the latter vein rather broad ; angle between the two veins relatively 
broader (Text-fig. 51) . . . . . fulvicornis sp. n. (p. 309) 

5 (3) Basal cell of fore wing extensively hairy. Funicle segments subequal in length, 

the third and fourth fuscous, the others yellowish; cava infuscate. In the 
only known female wings hyaline, but similar infumation as in male holotype, 
broadly appended to the veins, may be expected . varicornis sp. n. (p. 310) 

- Basal cell bare but sometimes bordered distally with a few hairs on basal and 

cubital fold. Funicle segments more distinctly decreasing in length, the 
distal ones never paler than the middle ones. Fore wing, if clouded, on the 
disc with an obliquely oval spot, not touching the veins .... 6 

6 (5) Flagellum all fuscous or hardly paler basally, very weakly clavate. 

dissimilis Walker (p. 313) 

- Funicle segments 1-4 usually paler brown, apex of flagellum blackish and 

slightly more clavate than above .... amoenus Forster (p. 312) 

Males 

1 Mid tibia at least partly infuscate and more or less broadened, flattened (Text- 

figs 56, 60, 61) ........... 2 

- Mid tibia all yellow and not distinctly enlarged ...... 5 

2 (1) Funicle segments alternately small and large (Text-fig. 47), the large ones partly 

brown, the small ones mainly pale yellow. Genae in facial view with a long 
comb of long white hairs (Text-fig. 46). Mid tibia with a broad external lobe 
(Text-fig. 56) ...... monospilus (Thomson) (p. 310) 

- Funicle segments hardly different in width, not alternating in colour. Genae 

with shorter hairs. Mid tibia more gradually broadened distally ... 3 

3 (2) Funicle segments 3 and 4 and clava distinctly infuscate, whilst funicle segments 

1-2 and 5-6 are yellow (Text-fig. 54). Flattening of mid tibia forms two 
waves on inner edge and one wave subapically on the outer edge (Text-fig. 61). 
Fore wing extensively infuscate in the middle, infuscation adhering broadly 
to marginal and stigmal vein ..... varicornis sp. n. (p. 310) 

- Distal funicle segments not paler-coloured than the middle ones. Mid tibia on 

inner side enlarged in a simple curve or almost straight. Fore wing infusca- 
tion, if present, less extensive and not adhering to the veins .... 4 

4 (3) Flagellum slender-filiform, along with pedicellus uniformly brownish testaceous 

(Text-fig. 55), distinctly darker than the pale yellow scape. Gena below with 
outstanding hairs which are about twice as long as the hairs on face below 
antennae. Mid tibia in dorsal view with inner side all blackish, more broadly 
flattened than the outer side. Eye longer than scapus. dissimilis Walker (p. 313) 



EUROPEAN PTEROMALIDAE 307 

- Flagellum subclavate, basally pale yellow, funicle segments 5 and 6 and clava fuscous. 

Gena below only with normal hairs which are subequal to those on lower face. 
Mid tibia in dorsal view with only the outer side flattened, the inner side 
usually with a pale line throughout (Text-fig. 60). Eye about as long as 
scapus ......... amoenus Forster (p. 312) 

5 (1) Head more strongly transverse than in the alternate 

Here probably the unknown male of S. polyspilus Graham 

- Head in dorsal view 1-78-1-96 times as broad as long, in facial view 1-21-1-29 

times as broad as high .......... 6 

6 (5) Fifth and sixth funicle segment subquadrate (Text-fig. 50) ; flagellum hardly 

longer than width of head, its segments uniformly flavous. Eye about 1 -25 
times as long as broad, about 1 -8 times as long as malar space. Scapus in 
lateral view hardly broader than the funicle. Thorax fairly arched dorsally. 
Apex of basal cell of fore wing more or less hairy. Possibly not associated 
with Phragmites ...... fulvicornis sp. n. (p. 319) 

Fifth and sixth funicle segment distinctly elongate; flagellum itself about i-i 
times as long as width of head, funicle segments partly and rather irregularly 
pale testaceous and partly brownish ; scapus in side view at least 1 -3 times as 
broad as the funicle. Thorax very weakly arched dorsally. Basal cell of 
fore wing bare. Associated with Phragmites . . peisonis (Erdos) (p. 317) 

Spaniopus polyspilus Graham 
(Text-figs 52, 57-59) 

[Polycelis conspersa (Walker); Thomson, 1878 : 143-144. Misidentification.] 
Spaniopus polyspilus Graham, 1956 : 251. Holotype $, Sweden: Stockholm (UZI, Lund) 
[examined]. [Proposed as new name for Polycelis conspersa (Walker) sensu Thomson, 1878.] 

This species is not conspecific with Spaniopus peisonis (Erdos) as Graham (1969) 
assumed, but a valid species the main characters of which are given in the key above. 

Biology still unknown. 

Known so far only from Sweden (Thomson's specimens) . 

Spaniopus peisonis (Erdos) 
(Text-figs 53, 62) 

[Polycelis monospila Thomson, 1878 : 145, ex parte (only $, not the lectotype).] 
Gyrinophagus peisonis Erdos, 1957 : 64, 6*9- Holotype $, Hungary: Vors (TM, Budapest) 
[examined]. 

I examined the Hungarian material including the holotype several years ago in 
Dr Erdos' collection. 

Biology. Parasite in the galls of Giraudiella inclusa (Erauenfeld) (Dipt., 
Cecidomyiidae) on Phragmites communis Trin. 

Distribution. Britain, Sweden, Czechoslovakia, Hungary. 



308 



Z. BOUCEK 




Figs 46-55. Spaniopus. 46-47. S. monospilns. 46, <J head; 47, $ antenna. 48-51. 
S. fulvicoriiis. 48, $ head; 49, 9 antenna; 50, $ antenna; 51, fore wing venation in ?. 
52. S. polyspilus, $ antenna. 53. 5. peisonis, $ head. 54. S. varicornis, o* antenna. 
55, 5. dissimilis. $ antenna. 



european pteromalidae 309 

Material examined. 

Type-data given in synonymy. 

Sweden : Smaland, 3 $ paralectotypes of P. monospila Thomson (Boheman) ; 
coll. Thomson, UZI, Lund ; Skane, Yddingen, 1 $ (Graham) ; Graham Coll. Czecho- 
slovakia: Bohemia, Bfehyne near Doksy, 3 $, 17.vii.1963 (Boucek); Kokofin, ex 
Giraudiella inclusa, 2$, vi.1958 (M. Skuhravd) ; Tyniste nad Orlici, 3^, 16 $, viii.1959, 
1 <$, 23.vii.1955 (Boucek); Sobeslav, 14 $, 22.vii.1955 (Hoffer); Moravia, Branisovice 
S. of Brno, 1 $, 29.V.1956 (Boucek); Slovakia, Turna nad Bodvou, 1 $, 23.iv.1952 
(Boucek) . 

Spaniopus fulvicornis sp. n. 

(Text-figs 48-51) 

$. Metallic green, on vertex and thoracic dorsum slightly merging with dark cupreous, 
gaster darker green; antenna testaceous, apically often slightly darker than basally, also 
pedicellus sometimes slightly infuscate; legs testaceous but coxae basally mainly dark and with 
metallic tinge, apex of tarsi infuscate. Wings hyaline but fore wing usually with fuscous 
cloud attached broadly to marginal and stigmal vein ; venation testaceous. Length 1 -9-2 -g 
(holotype 2-6) mm. 

Head dorsally fully twice as broad as long (49 : 24), 1-24 times as broad as mesoscutum and 
1-49 times as broad as pronotum; temple about one-third length of eye in dorsal view. POL 
to OOL as 4 : 3. Relative measurements: breadth of head 49, height 40, breadth of frons 33, 
eye 24 : 17, malar space 13, width of mouth 23, scapus length 205, flagellum plus pedicellus 43, 
i.e., o-88 the breadth of head. Left mandible 3, right 4 teeth. Clypeus and lower face densely 
radiately striate, lower margin of clypeus subemarginate. Scapus not quite reaching vertex 
level, slender, bent (Text-fig. 49). Pedicellus dorsally 2-2-1 times as long as broad; second 
anellus only slightly transverse (as in S. polyspilus) ; funicle segments distinctly decreasing in 
length and increasing in width, the first in some views narrower than pedicellus and about 1 -5 
times as long as broad, the sixth in lateral view 1-5 times as broad as pedicellus and 1-5 times 
as broad as long; clava barely longer than two preceding segments combined. 

Length of thorax measured from anterior edge of collar 1 -53 times the breadth of mesoscutum. 
Collar moderately sloping, medially 0-22 the length of mesoscutum. Scutellum with rather 
deep reticulation-puncturation. Propodeum medially two-thirds the length of scutellum, 
nucha taking up posterior 3/7; median area rather dull, irregularly reticulate-punctured, as 
broad as median length of propodeum. Plicae distinct, almost regularly arcuate; part beyond 
plicae densely clothed with white hairs, except area around spiracle. Upper mesepimeron 
smooth. Hind femur 45 times as long as broad. Fore wing: relative lengths of veins: 
marginal 22, postmarginal 13, stigmal vein 11, the latter at a relative^ wide angle (Text-fig. 
51). Basal cell with adjoining folds bare, basal fold sometimes with 1-2 hairs; lower surface 
of costal cell basally with single hair-line. 

Gaster barely as long as head plus thorax combined, ovate-acuminate, itself 1 -66-I -85 times 
as long as broad, sides of sixth tergite converging at about 6o°. 

<J. Similar to female in colour and in form of thorax. Antennae and legs yellowish testaceous. 
Scapus slightly exceeding vertex level, as long as eye (in one specimen slightly shorter than eye). 
Flagellum (Text-fig. 50) plus pedicellus about i-i times as long as head width; pedicellus 
dorsally i-8 times as long as broad; flagellum feebly clavate, unicolorous, first funicle segment 
about 1-5 times as long as broad, the sixth subquadrate; clava about 2-5 times as long as broad, 
subacuminate. Genae without outstanding hairs. Mid tibia simple and not infuscate. Fore 
wing subhyaline, with basal cell slightly hairy in distal part; marginal vein about i-8 times 
(1 75-1 -89 times) as long as the stigmal. Gaster slightly longer than half length of thorax. 
Length of body 1 -5-1 -6 mm. 



310 Z. BOUCEK 

Biology unknown. Probably a grass-dweller. 

Holotype $. Czechoslovakia: Bohemia, Sedlo Hill near Litomefice, 6.viii.ig64 
(Boucek) ; presented to BMNH, London. 

Paratypes. Czechoslovakia: Bohemia, Mt. Decinsky Sneznik, 2 $, 1 £, 2y.vii. 
IQ56 (Boucek) ; Bela near Deem, 3 $, 2 $ (one of them allotype), 20.viii.1956 (Boucek) ; 
Bfehyne near Doksy, 1 $, 21.vii.1963 (Strejcek); Jedlova Mt., near Rumburk, 1 $, 
8.V.1960 (Boucek); Slovakia, Smokovec, High Tatra Mts, 1 $, 21.viii.1958 (Boucek). 
Paratypes partly in NM, Prague, partly in Boucek Collection. 

Spaniopus monospilus (Thomson) 
(Text-figs 46, 47, 56) 

Polycelis monospila Thomson, 1878 : 145. Lectotype o*> Sweden: Kinekulle (UZI, Lund) 
[examined] . 

Polyscelis Websteri Ashmead, 1894: 52-53, ?q\ Holotype $, U.S.A.: Indiana, Lafayette 
(USNM, Washington) [examined]. Syn. n. 

P. websteri is a synonym of S. monospilus as Graham (1969 : 706) has already 
suggested. In October 1970 I showed the very distinctive lectotype male of mono- 
spilus to Dr Burks and he agreed with the synonymy. More recently he kindly sent 
to me for examination the holotype of websteri. Because the females referred to 
5. monospilus by Thomson proved to belong to S. peisonis (see under that species), 
the holotype of websteri is the only known female of the present species. Its 
characters are included in the key above. 

Biology not known with certainty. P. websteri was recorded from a Cynipid 
gall on Lactuca canadensis (Peck, 1963), which seems doubtful (Dr B. D. Burks, 
personal communication) . 

Distribution: Sweden (only 2 males); U.S.A. 

Spaniopus varicornis sp. n. 

(Text-figs 54, 61) 

$. Mainly metallic green, but vertex dark purple to violaceous or bronzy, thoracic dorsum 
with dull cupreous to bronzy tinge, gaster bluish green with bright green base. Antenna 
testaceous with funicle segments 3 and 4 fuscous; weakly infuscate, mainly dorsally, are also 
pedicellus, second and fifth funicle segment; clava fuscous. Wings hyaline, venation testaceous. 
Length 2 mm. 

Head dorsally twice as broad as long, 1 -26 times as broad as mesoscutum 1 -53 times as broad 
as pronotum; temple one-third the length of eye. POL 1-4 times the OOL In facial view 
head 1 -26 times as broad as high, with genae strongly converging (at about 120 ). Lower face 
very distinctly radiately striate, lower margin of clypeus shallowly depressed, subemarginate. 
Scapus reaching to level with anterior edge of median ocellus; pedicellus dorsally about i-8 
times as long as broad; flagellum plus pedicellus as long as 0-9 the breadth of head; both anelli 
together about as long as broad; flagellum in lateral view slightly clavate, clothed with semi- 
erect hairs almost half as long as segments; all funicle segments subequal in lengths, in lateral 



EUROPEAN PTEROMALIDAE 



3ii 



view the first slightly elongate, the sixth slightly transverse; clava fully twice as long as broad. 

Thorax almost as in male. Legs normal, not very slender, hind femur about 4-3 times as 
long as broad, hind tarsus 0-7 the length of tibia. Fore wing with basal cell extensively hairy. 
Relative lengths of veins: marginal 12-5, postmarginal 12, stigmal vein 11. 

Gaster ovate-pointed, slightly longer than thorax, itself 1-62 times as long as broad; in 
dorsal view sides of sixth tergite converging at angle of about 8o°. 

<J. Bright green with weak brassy reflections on frons and thoracic dorsum. Antenna 
pale yellow, but funicle segments 3 and 4 fuscous; apex of scapus, pedicellus and clava except 
apex also infuscate (Text-fig. 54). Tegulae and legs including coxae pale testaceous, but mid 




Figs 56-62. Spaniopus. 56. S. monospihts, mid tibia in <J. 57-59- 5. polyspilus. 
57, fore wing in $; 58 & 59, $ head in facial and dorsal view. 60. S. amoenus, mid tibia 
of (J- 61. S. varicornis, mid tibia of <J. 62. S. peisonis, fore wing of $ with well 
developed markings. 



312 Z. BOUCEK 

tibia fuscous except basally (Text-fig. 61). Fore wing with broad brownish cloud attached to 
marginal and stigmal vein. Length i-6 mm. 

Head dorsally 1-91 times as broad as long, eye 3-3 times as long as temple; POL 1-5 times the 
OOL. In facial view head 1 -24 times as broad as high. Lower face more distinctly, clypeus 
more finely and shallowly, radiately striate, in middle below smooth; lower margin sube- 
marginate. Mandibles: left 3, right 4 teeth. Genae arched, strongly converging, below with 
group of erect white hairs of medium length. Relative measurements: breadth of head 34, 
frons 22, eye 17-2 : 13, malar space 7, width of mouth 15-4, length of scape 15-5, flagellum plus 
pedicellus about 46, i.e., 1-36 times the breadth of head. Pedicellus dorsally 17 times as long 
as broad ; both anelli transverse, together hardly as long as broad ; funicle segments (Text-fig. 
54) subequal in length, hardly increasing in breadth, the basal ones inconspicuously longer 
than broad, the distal ones subquadrate; clava slightly shorter than three preceding segments 
together; flagellum with semidistant hairs slightly shorter than segments. 

Pronotum moderately narrower than mesoscutum, rather deeply emarginate posteriorly, 
collar anteriorly angulate, in middle about 1/6 the length of mesoscutum. The latter hardly 
more than 1-5 times as broad as long; notauli fading out in the middle. Propodeum 0-78 the 
length of scutellum, median area about 1-1 times as broad as long, anteriorly less coarsely 
more than 1-5 times as broad as long; notauli fading out in the middle. Propodeum 0-78 the 
reticulate than on nucha. Plicae distinct but not sharp; small oval spiracle nearly two 
diameters from metanotal margin; lateral part of propodeum beyond spiracle and beyond 
posterior half of plica densely hairy. Legs rather strong: femora slightly thickened, also fore 
tibia which is in side view only 5-3 times as long as broad. Mid tibia (Text-fig. 61) flattened, 
in dorsal view externally enlarged only in distal half, internally both in basal and distal halves, 
in outline there forming double curve. Mid femur 4 times as long as broad. Fore wing with 
basal cell hairy all over. Relative lengths of veins: marginal 15, postmarginal 12, stigmal 9-5. 

Gaster subrotund, broader and shorter than thorax. Petiole conspicuous, smooth, in middle 
slightly broader than long, sides anteriorly parallel. First tergite covering more than half of 
gaster, its hind margin arcuate. 

Biology not known. 

Holotype $. Czechoslovakia: Slovakia, Smokovec in High Tatra Mts, 2o,.viii. 
1958 {Boucek) ; in Boucek Collection. 

Paratype. Czechoslovakia: Bohemia, Tyniste nad Orlici, 1 $ (allotype), 
12. ix. 1959 (Hoffer); in Graham Collection. 

The absence of the fore wing infumation in the allotype may be due to the sub- 
teneral condition of the specimen. 

Spaniopus amoenus Forster 

(Text-fig. 60) 
Spaniopus amoenus Forster, 1856 : 56. Type o*. Germany: ? Aachen (Plost). 

The type-material is probably lost but the short description fits the fresh material 
well. The female was not then known. It is extremely similar to that of Spaniopus 
dissimilis Walker and except for the colour of the antenna in most specimens 
(which, however, does not always seem to be reliable) I cannot find any additional 
character. Various parts of the body which are likely to yield some difference were 
measured and show a rather wide range of variation. Length of scape in relation 



EUROPEAN PTEROMALIDAE 313 

to the long eye diameter varies from 0-91-1-0 (: 1), while in 5. dissimilis the same 
relation shows figures between 0-85 and 0-93. Similar overlap has been found in 
the relative length of malar space and of the veins in the fore wing. Length of body 
1-8-2-4 mm - 

Biology not known, but the species seems to be associated with grasses in xero- 
dermic habitats. 

Distribution. France, W. and E. Germany, Czechoslovakia, Hungary. 

Material examined. 

France: Finisterre, Huelgoat, 2 $, 29.V.1954 (/. F. Perkins), in BMNH. E. 
Germany: Dresden district, 1 J, 16.ix.1965 (Strejcek). Czechoslovakia: Bohemia, 
Dzban Hill, 3 $, 2 £, 4.ix.i966 (Boudek); Velky Vfestov, 1 <$, viii.1961 (Boucek); 
Slovakia, Slanec, incl. Lake Izra and Helmec Valley, 6 $, 1 <$, 3.-6. viii. 1954 (Boudek 
& Dlabola). Hungary: Mecsek Hills, Misina, sifting under heath, about 80 $ and 
several^, 24.x. 1953 (Kaszab); Budapest-Hiivosvolgy, 1 $, 2 <$, vi.-ix.1927 {Biro), 
the Hungarian material mostly in TM, Budapest. 



Spaniopus dissimilis Walker 
(Text-fig. 55) 

Spaniopus dissimilis Walker, 1833 : 466. Holotype <$, Britain: near London (BMNH) 

[examined] . 
Spaniopus elegans Forster, 1856 : 56. Holotype <J, W. Germany: (?) Aachen (?lost). Syn. n. 
Polyscelis modestus Gahan, 1922 : 11-12, $q*- Holotype $, U.S.A.: Pennsylvania, Hannover 

(USNM, Washington). 

Graham (1969 : 705) is probably right in regarding P. modestus as a synonym of 
S. dissimilis, although the figure 16D in Gahan, 1933, shows the mid tibia of the male 
a little too slender. From the material examined I conclude that also 5. elegans 
Forster, although its type seems to be lost, must be the same species. The flagellum 
of the male is usually brownish, but sometimes paler, yellowish, as described for 
S. elegans. 

Biology. Solitary ectoparasite of Mayetiola destructor (Say) (Dipt., Cecido- 
myiidae) in grass stems, including wheat ; mostly primary, rarely secondary (Gahan, 
1922, 1933). According to my experience, unlike the closely related S. amoenus, 
S. dissimilis is not associated with xerothermic habitats. 

Distribution. Ireland, Britain, Sweden, Czechoslovakia; Canada, U.S.A. 

New records. 

Britain: Ross-shire, Black Isle, 1 $, vii.1951 (N. Hussey). Sweden: Skane, 
Fjellfota sjo, 1 $, 1 $, 31.vii.1938 (/. F. Perkins) ; Degaberga, 1 <J, 8.vii.i938 (D. M. S. 
P. & J. F. P.) ; both in BMNH. Czechoslovakia: Bohemia: Bela near Decin, 1 <£, 
14. vi. 1957 (Boucek) ; Bfehyne near Doksy, 2 $, 1 <$, 12.vii.1959 (Boucek) ; Tyniste nad 
Orlici, 1 <J, 23.vii.1955 (Boucek); partly in NM, Prague. 



3M Z. BOUCEK 

REFERENCES 

Ashmead, W. H. 1894. Descriptions of thirteen new parasitic Hymenoptera, bred by 

Prof. F. M. Webster. /. Cincinn. Soc. nat. Hist. 17 : 45-55, 2 pis. 
Boucek, Z. 1958. Eine Cleonyminen-Studie ; Bestimmungstabelle der Gattungen mit 

Beschreibungen und Notizen, eingeschlossen einige Eupelmidae (Hym. Chalcidoidea). 

Sb. ent. Odd. ndr. Mus. Praze 32 : 353-386, 40 figs. 
1965. Some interesting records of Chalcid flies from Great Britain, with the description 

of Bugacia classeyi n. sp. (Hymenoptera: Pteromalidae) . Entomologist's Gaz. 16 : 83-86, 

4 figs. 

1970. Contribution to the knowledge of Italian Chalcidoidea (Hym.) based mainly on a 

study at the Institute of Entomology in Turin, with descriptions of some new European 
species. Mem. Soc. ent. ital. 49 : 35-102, 16 figs. 

Erdos, J. 1953. Pteromalidae hungaricae novae. Acta biol. hung. 4 : 221-247, ZI n S s - 

1957- Recentiores observationes entomocoenologicae in Phragmite communi Trin. [In 

Hungarian and Latin]. Allat. Kozl. 46 : 49-65, 7 figs. 

Fabricius, J. C. 1798. Supplementum Entomologiae systematicae. 572 pp. Copenhagen. 

Forster, A. 1856. Hymenopterologische Studien. 2. Chalcidiciae und Proctotrupii . 152 pp. 

Aachen. 
Gahan, A. B. 1922. Descriptions of miscellaneous new reared parasitic Hymenoptera. 

Proc. U.S. natn. Mus. 61 : 1-24, 1 pi. 
■ 1933- The Serphoid and Chalcidoid parasites of the Hessian fly. Misc. Pubis U.S. Dep. 

Agric. 174 : 1-147, 32 figs. 
Graham, M. W. R. de V. 1956. A revision of the Walker types of Pteromalidae (Hym., 

Chalcidoidea). Part 2 (including descriptions of new genera and species). Entomologist's 

mon. Mag. 92 : 246-263, 6 figs. 

1969. The Pteromalidae of north-western Europe (Hymenoptera: Chalcidoidea). Bull. 

By. Mus. nat. Hist. (Ent.) Suppl. 16 : 1-908. 

Kerrich, G. J. & Graham, M. W. R. de V. 1957. Systematic notes on British and Swedish 
Cleonymidae, with description of a new genus (Hym., Chalcidoidea). Trans. Soc. Brit. 
Ent. 12 : 265-311, 2 pis, 24 figs. 

Kryger, J. P. 1943. The Chalcid subfamily Eunotinae. Ent. Meddr 23 : 66-8 1, 6 figs. 

Kurdjumov, N. V. 1912. Six new species of Chalcid flies parasitic upon Eriococcus greeni 
Newstead. Russk. ent. Obozr. 12 : 329-335, 8 figs. 

Latreille, P. A. 1809. Genera Crustaceorum et Insectorum. 4, 399 pp. Paris & Strasbourg. 

Masi, L. 1907. Contribuzioni alia conoscenza dei Calcididi italiani. Boll. Lab. Zool. gen. 

agr. R. Scuola Agric. Portici 1 : 231-295, 47 figs. 
1928. Diagnosi di una nuova specie di Eunotus (Hymen. Chalcididae). Boll. Soc. ent. 

ital. 60 : 128. 

1931. Contribute alia sistematica degli Eunotini (Hym. Chalc). Eos, Madr. 7 : 411-459, 

6 figs. 

Nikolskaya, M. N. & Kyao, N. N. 1954- [Chalcid fauna of the middle course of the river 

Ural and its economic importance.] [In Russian]. Trudy zool. Inst. Leningr. 16 : 404-416, 

6 figs. 
Peck, O. 1963. A Catalogue of the Nearctic Chalcidoidea (Insecta: Hymenoptera). Can. 

Ent., Suppl. 30. 1092 pp. 
Peck, O., Boucek, Z. & Hoffer, A. 1964. Keys to the Chalcidoidea of Czechoslovakia 

(Insecta: Hymenoptera). Mem. ent. Soc. Can. no. 34. 121 pp., 289 figs. 
Ratzeburg, J. T. C. 1852. Die Ichneumonen der Forstinsecten in forstlicher und entomologischer 

Beziehung, ein Anhang zur Abbildung und Besckreibung der Forstinsecten. 3. vi-xviii + 272 

pp., 3 tables. Berlin. 



EUROPEAN PTEROMALIDAE 



315 



Swederus, N. S. 1795. Beskrifning pi et nytt genus Pteromalus ibland Insecterna, hoerande 
til Hymenoptera. K. svenska VetenskAkad. Handl. 16 : 201-205, 216-222. 

Thomson, C. G. 1878. Hymenoptera Scandinaviae. 5. Pteromalus (Svederus) continuatio. 
307 pp., 1 pi. Lund. 

Walker, F. 1833. Monographia Chalcidum. Ent. Mag. 1 : 367-384, 455-466. 

1834. Monographia Chalciditum. Ent. Mag. 2 : 148-179, 286-309, 340-369 

1837. Monographia Chalciditum. Ent. Mag. 4 : 349-358. 

Westwood, J. O. 1839. Synopsis of the genera of British insects. Pp. 49-80. London. 



PARASITE INDEX 



acerina, Susteraia, 289 
acutus, Eunotus, 285, 275, 277 
amoenus, Spaniopus, 312, 306, 307 
antshar, Eunotus, 282 
aquisgranensis, Eunotus, 287 
areolatus, Eunotus, 282, 275, 276 

balcanicus, Cleonymus, 270, 269 
brevior, Strejcekia, 297, 296 
brevis, Cleonymus, 272, 269 

Cecidostiba, 299 
Cleonymus, 269 
congrua, Peridesmia, 305 
conspersa (Polycelis), 307 
crassicornis, Habrocytus, 313 
cretaceus, Eunotus, 281, 275, 276 
cupreus, Rhizomalus, 300 

depressus, Cleonymus, 273 
discus, Peridesmia, 305 
dissimilis, Spaniopus, 313, 306 
Dorcatomophaga, 294 

elegans, Spaniopus, 313 
elegans, Strejcekia, 297, 296 
Eunotellus, 274 
Eunotus, 274 

festucae, Eunotus, 281 

fulvicornis, Spaniopus, 309, 306, 307 

Habrocytus, 303 

Hobbya, 299 

hofferi, Eunotus, 277, 275, 276 

kocoureki, Eunotus, 279, 275, 276 
Ksenoplata, 289 

laticornis, Cleonymus, 270, 273 



Megapelte, 274 
merceti, Eunotus, 286, 276 
modestus (Polyscelis) , 313 
monospilus, Spaniopus, 310, 306 
montana, Peridesmia, 303, 304 

Nephelomalus, 299 

nigriclavis, Eunotus, 286, 275, 276 

Nodisoplata, 289 

obscurus, Cleonymus, 270, 275 
obscurus, Eunotus, 284, 275 

paludicola, Pteromalus, 302, 303 

parvulus, Eunotus, 287, 274, 277 

Pegopus, 300 

peisonis, Spaniopus, 307, 305 

Peridesmia, 303 

polyspilus, Spaniopus, 307, 305 

Pteromalus, 302 

rara, Veltrusia, 294 
Rhizomalus, 298, 299, 300 
rujanensis, Semiotellus, 291 

Semiotellus, 290 
Semiotus, 290 
Spaniopus, 305 
Strejcekia, 295, 292, 294 
subcyaneus, Eunotus, 282 
Susteraia, 287, 289 

Tritypus, 292, 274 

varicornis, Spaniopus, 310, 306 
Veltrusia, 292, 294 
venustus, Pteromalus, 303 
vopiscus, Pteromalus, 303 

websteri (Polyscelis), 310 



Z. Boucek 

Commonwealth Institute of Entomology 

c/o British Museum (Natural History) 

Cromwell Road 

London, SW7 5BD 



3 C o Cl 



^ 



h 



A LIST OF SUPPLEMENTS 

TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp.177: 
18 plates, 270 text-figures. August, 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 : 500 text-figures. September, 

1965- £3-25- 

5. Ahmad, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156: 
475 text-figures. November, 1965. (out of print) £2.15. 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. 

£3-15- 

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the 
world species of Chora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 
text-figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera) . Pp. 509. £8.50. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322: 348 text-figures. August, 1967. £8. 

11. Mound, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp.172: 
82 text-figures. May, 1968. £4. 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. 
November, 1968. £5. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North-Western Europe 
(Hymenoptera : Chalcidoidea) . Pp. 908: 686 text-figures. November, 1969. 

£i9- 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 

68 plates, 15 text-figures. October, 1971. £12. 



j>(S»)o Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 



- 

53 



[ 1 6 JAN (973 

THE ANT GENERA OF WEST AmiCA: 
A SYNONYMIC SYNOPSIS 

WITH KEYS 
(HYMENOPTERA : FORMICIDAE) 



B. BOLTON 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 27 No. 6 

LONDON : 1973 



THE ANT GENERA OF WEST AFRIC 
A SYNONYMIC SYNOPSIS WITH KE 
(HYMENOPTER A : FORMICIDAE) 



4* 




BY 



BARRY BOLTON 



Pp. 317-368; 1 Text-figure 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol.27 No. 6 

LONDON: 1973 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 27 No. 6 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Ent.). 



Trustees of the British Museum (Natural History), 1973 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



Issued 18 January, 1973 Price £2.60 



THE ANT GENERA OF WEST AFRICA: 

A SYNONYMIC SYNOPSIS WITH KEYS 

(HYMENOPTERA : FORMICIDAE) 

By B. BOLTON 



CONTENTS 

Synopsis ........ 

Introduction ....... 

Acknowledgements ...... 

Definitions of some characters used in the keys 
Key to the subfamilies of Formicidae in West Africa 
Keys to the genera 

Subfamily Ponerinae 

Subfamily Cerapachyinae 

Subfamily Myrmicinae 

Subfamily Dorylinae 

Subfamily Leptanillinae 

Subfamily Pseudomyrmecinae 

Subfamily Dolichoderinae 

Subfamily Formicinae 
Synonymic synopsis and notes on the 
References . 
Index .... 



Page 
319 
319 
320 
320 
321 
323 
323 
325 
325 
328 
328 
329 
329 
329 
330 
364 
366 



SYNOPSIS 

Keys are given to the subfamilies and genera of ants present in West Africa. A synopsis of 
the known generic synonymy is presented along with diagnostic notes on the genera. 

INTRODUCTION 

West Africa, in the sense of the present study, includes the territories of Senegal, 
Gambia, Portugese Guinea, Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, 
Togo, Dahomey, Nigeria, Upper Volta, and the southern portions of Mali and 
Niger (below 15 ° N.). 

Two distinct types of vegetation occur in the above states, forest and savannah. 
The forest zone forms a belt of varying width running parallel to the coast, some- 
times coming very close to the sea but often separated from the ocean by a coastal 
plain. The forest zone is not represented in the inland states of Upper Volta, 
Mali and Niger. North of the forest zone is the inland savannah belt which 
grades into the Sahara Desert. In places, such as the Dahomey forest gap, the 
savannah reaches to the coast and the ant fauna of the coastal plains bears a greater 
resemblance to the savannah than to the forest population. 

As defined above, West Africa is bounded on the west and south by the Atlantic 
Ocean, on the north by the steppe region which grades into the Sahara Desert, 
and on the east by the mountain ranges of Cameroun. 



320 B. BOLTON 

The majority of ant genera in West Africa contain species in both the forest and 
savannah zones, but some genera are confined to the forest (e.g. Psalidomyrex 
E. Andre) whilst others are restricted to the savannah and the coastal plains (e.g. 
Messor Forel, Cataglyphis Forster). 

Wheeler (1922) included keys to the world genera of ants and a check list of the 
ants of the Ethiopian Region in the monumental 'Ants of the Belgian Congo'. These 
keys are now considerably outdated due to revisions, synonymies and descriptions 
of new genera carried out since the publications of this study, and are now unwork- 
able in many places. 

The first section of the present paper gives keys to the subfamilies and genera of 
ants recognized at present from West Africa, and includes some pantropical tramp 
ants such as Trichoscapa Emery, which have not yet been reported but can be 
expected to be found in the region. In the second section the genera are arranged 
under their appropriate subfamilial and tribal groupings, and the known synonyms 
up to the time of writing are included, as are short diagnostic notes on the genera. 
An indication of nesting sites and habits are given where these are known. 



ACKNOWLEDGEMENTS 

I am grateful to the directors of the Cocoa Research Institutes of Nigeria and 
Ghana for providing equipment and space during the course of this survey, and to 
the International Office of Cocoa and Chocolate for financial support. 

I would also like to express my thanks to Messrs D. Leston and P. M. Room of the 
University of Ghana, Legon, and to Mr C. A. Collingwood of the International 
Capsid Research Team for access to their personal collections of West African ants. 



DEFINITIONS OF SOME CHARACTERS USED IN THE KEYS 

(See also Text-fig. 1) 

Acidopore: a circular or subcircular orifice formed by the apex of the hypopygium 

in the subfamily Formicinae, the orifice of the acid-producing glands. Usually 

the acidopore is visible at the apex of the gaster, sometimes borne on a conical 

projection of the hypopygium but occasionally concealed by the pygidium when 

not in use. (Hung & Brown, 1966). 
Alitrunk (= mesosoma) : the thorax proper plus the propodeum (first true abdominal 

segment) to which it is fused in the higher Hymenoptera. The 'thorax' of older 

authors. 
Antennal Scrobe: a longitudinal depression in the side of the head, either above or 

below the eye, which can accommodate the scape or the whole of the antenna. 
Frontal Carinae: a pair of longitudinal ridges situated mediodorsally on the head 

behind the clypeus. Usually the frontal carinae are expanded laterally into 

lobes which cover the antennal insertions. 
Funiculus (= flagellum): collective term for all the antennal segments distal to the 

elongated first segment or scape. 



ANT GENERA OF WEST AFRICA 321 

Gaster: the four or five visible remaining abdominal segments situated, behind the 
separated pedicellar segments, and forming the apparent 'abdomen'. 

Hypopygium: the last visible gastral sternite (bearing the acidopore in the sub- 
family Formicinae). In reality the sternite of the seventh abdominal segment. 

Palp Formula: the number of segments in the maxillary and labial palpi, always 
expressed in the order maxillary, labial, (e.g. palp formula 6,4.) 

Pedicel : the one or two separated abdominal segments between the alitrunk (apparent 
thorax) and gaster (apparent abdomen). The first segment is termed the petiole, 
the second when present the postpetiole. In reality the second and third (when 
present) true abdominal segments. 

Petiole and Postpetiole : see Pedicel, above. 

Propodeum (= epinotum): the first true abdominal segment, fused to the thorax 
proper to form the alitrunk. 

Psammophore : a basket-like arrangement of long setae found on the gular surface 
of the head in deserticolous ants and used to transport grains of soil. 

Pygidium: the last visible gastral tergite. In reality the tergite of the seventh 
true abdominal segment. 

Scape: the elongate first segment of the antenna. 

Keys to the Subfamilies of FORMICIDAE in West Africa 
(based on worker caste) 

1 Pedicel of a single segment, usually with a narrow connection to the gaster so that 

the petiole has a distinct posterior face. Rarely the petiole is reduced or very 
broadly attached to the gaster ......... 2 

- Pedicel of two segments, usually with the postpetiole distinctly separated from the 

gaster. Rarely the postpetiole is broadly attached to the gaster, but in this case 

the antennae are only 6-segmented ........ 6 

2 Eyes absent. Clypeus reduced so that the antennal insertions are very close to the 

anterior margin of the head. Frontal carinae raised, leaving the condylar bulbs of 
the antennae visible in dorsal view. Pygidium impressed, armed with a spine or tooth 
at each side posteriorly. Promesonotal suture distinct, mobile. Lateral alitrunk 
in large and medium workers with a distinct, deep longitudinal impression below 
the propodeal spiracle. Sting reduced, non-functional. Polymorphic. (Driver 
Ants) DORYLINAE (p. 328) 

- Without all the above characters. Eyes usually present; clypeus developed so that 

the antennal insertions are some distance behind the anterior margin of the head. 
Frontal carinae usually at least partially cover antennal insertions ; if not, then the 
dorsum of the alitrunk is without sutures. Pygidium rarely impressed; alitrunk 
laterally without a deep impression below the propodeal spiracle .... 3 

3 Sting well developed, functional. (Usually visible in alcohol-killed or mounted 

specimens) ............. 4 

- Sting vestigial or absent, never visible ......... 5 

4 Pygidium impressed, armed laterally or posteriorly with a row of short denticles or 

spines (which may be difficult to see due to setal development). Genae carinate. 
Frontal carinae fail to cover condylar bulbs of antennae. Alitrunk dorsally com- 
pletely devoid of sutures, or sutures represented by weak impressions at most 

CERAPACHYINAE (p. 325) 

- Pygidium never impressed, never armed with spines or denticles. Genae usually not 

carinate; frontal carinae usually cover condylar bulbs of antennae. Alitrunk 



322 



B. BOLTON 



dorsally usually with at least one suture; if sutures absent, then the second gastral 
segment is strongly vaulted ...... PONERINAE (p. 323) 

Apex of gaster with a circular acidopore formed from the hypopygium, this structure 
often projecting as a nozzle and fringed with setae. Occasionally the orifice of 
the acidopore is hidden by a projection of the pygidium, in which case the pronotum, 
petiole or both armed with spines. Petiole usually a scale or node, rarely reduced 

FORMICINAE (p. 329) 

Acidopore absent, the gaster terminating in a transverse slit bounded by the pygidium 
and hypopygium. Petiole usually reduced and overhung by the first gastral seg- 
ment, rendering the petiole invisible in dorsal views. DOLICHODERINAE (p. 329) 

Pygidium impressed and armed laterally or posteriorly with a row of short spines or 
denticles (which may be difficult to see due to setal development) 

CERAPACHYINAE (p. 325) 

Pygidium not impressed, not armed with teeth or denticles. ..... 7 

Frontal carinae vertical, failing to cover the antennal insertions ; eyes absent . . 8 

Frontal carinae not vertical, totally or partially covering the antennal insertions ; eyes 

usually present ............ 9 

Antennae 10-segmented; genae carinate. Larger ants with head width greater than 

0-25 mm DORYLINAE (p. 328) 

Antennae 12-segmented; genae not carinate. Minute ants with head width less than 

0-25 mm LEPTANILLINAE (p. 328) 



claw — 




mandible 
. gena 



funiculus 

pronotum 

mesonotum 

propodeum 

petiole 



gaster 



Fig. 1. Worker of Brachyponera senaarensis (Mayr), to illustrate some terms used in the key. 



ANT GENERA OF WEST AFRICA 323 

Clypeus projecting back between the frontal carinae. Tibial spurs of middle and 
hind legs usually simple or absent; if pectinate a psammophore is present on the 
gular surface of the head. Antennae 4- to 12-segmented. Eyes usually medium 
to small in size, ocelli absent. Claws simple . . MYRMICINAE (p. 325) 

Clypeus not projecting back between frontal carinae. Tibial spurs of middle and hind 
legs pectinate. Psammophore absent. Antennae always 12-segmented. Eyes 
usually large, ocelli often present. Claws usually toothed 

PSEUDOMYRMECINAE (p. 329) 

Keys to the Genera 

(based on worker caste) 

Subfamily PONERINAE 

Petiole broadly attached to first gastral segment, without a free posterior face. 

Mandibles elongate and narrow, dentate down the entire inner margin . . 2 

Petiolar-gastral junction narrow, petiole usually with a distinct free posterior face. 

Mandibles not as above; if elongate they are either not dentate down the entire 

inner margin or are not narrow ......... 3 

Mandibles pointed at apex, not as long as head; tooth row on inner margin single. 

Sculpture fine, spatulate hairs absent . . AMBLYOPONE Erichson (p. 330) 

Mandibles blunt at apex, very long (longer than head), with a double tooth-row on 

the inner margin. Sculpture coarse, spatulate hairs present 

MYSTRIUM Roger (p. 331) 
Tergite of second gastral segment strongly vaulted so that the remaining segments 

point anteriorly. Alitrunk devoid of sutures ....... 4 

Tergite of second gastral segment not strongly vaulted, remaining segments directed 

posteriorly. Alitrunk usually with at least one suture visible dorsally . . 5 

Mandibles edentate, overhung by the projecting clypeus. Apical funicular segment 

bulbous DISCOTHYREA Roger (p. 332) 

Mandibles with three teeth, not overhung by clypeus. Apical funicular segment 

only moderately enlarged . . . PROCERATIUM Roger (p 333) 

Mandibles linear, inserted in the middle of the anterior margin of the head, with an 

apical armament of three teeth arranged in a vertical series .... 6 

Mandibles inserted at sides of anterior margin of head, not armed apically with a 

vertical series of three teeth .......... 7 

Dorsalmost tooth of apical mandibular series acute . ANOCHETUS Mayr (p. 333) 

Dorsalmost tooth of apical mandibular series truncated. 

ODONTOMACHUS Latreille (p. 333) 
Claws pectinate (incompletely so in some species) . 

Clypeus carinate, projecting as a lobe or point antero-medially. Mandibles 

inserted at extreme corners of anterior margin of head, either edentate or nearly so 

LEPTOGENYS Roger (p. 337) 
Claws simple or with a tooth, never pectinate ....... 8 

Middle tibiae and tarsi with abundant downcurved spines and stiff hairs on the 

outer surface, giving a brush-like appearance. Tibial spurs of middle legs both 

small and simple, hind tibiae with a large pectinate and a small simple spur. 

Eyes absent CENTROMYRMEX Mayr (p. 335) 

Middle tibiae and tarsi not as above. Tibial spurs of middle and hind legs similarly 

developed. Eyes usually present ......... 9 

Mandibles elongate, linear, somewhat curved . . . . . . . 10 

Mandibles triangular or subtriangular . . . . . . . . 12 

Mandibles blunt and edentate apically, the inner margin with not more than two 

blunt teeth. Mandibular articulation associated with a marked semicircular 

excavation of the dorsal anterior margin of the head in front of the eye . . 11 



324 B. BOLTON 

- Mandibles dentate apically, their inner margins retaining feeble traces of teeth. 

Mandibular articulation not associated with a semicircular excavation of the 
anterior margin of the head .... PROMYOPIAS Santschi (p. 339) 

1 1 Mandibles with distal half swollen, the inner margin without teeth 

CACOPONE Santschi (p. 335) 

- Mandibles with distal half tapering, not swollen; inner margin with one or two 

blunt teeth PLECTROCTENA F. Smith (p. 339) 

12 Clypeus fused to frontal carinae and forming a plate which projects forwards above 

the mandibles. Antennae inserted close to the anterior margin of this plate, their 
condylar bulbs exposed .... PROBOLOMYRMEX Mayr (p. 332) 

- Clypeus and frontal carinae not modified as above; condylar bulbs of antennae 

covered by frontal carinae . . . . . . . . . .13 

13 Gaster without a constriction between the first and second segments ... 14 

- Gaster with a constriction between the first and second segments. . . . 15 

14 Petiole armed with a comb of five teeth which recurve over the gaster 

PHRYNOPONERA Wheeler (p. 338) 

- Petiole a narrow, unarmed, erect scale . . ASPHINCTOPONE Santschi (p. 334) 

15 Pretarsal claws armed with a tooth either medially or near base . . . . 16 

- Pretarsal claws simple, without teeth . . . . . ... 18 

16 Gena with a carina running longitudinally between the eye and the mandibular 

articulation. Claws toothed near the base . . MEGAPONERA Mayr (p. 337) 

- Gena not carinate. Claws with a median tooth . . . . . . . 17 

17 Tibiae of middle and hind legs each with two pectinate spurs. Sculpture of fine 

dense shagreening with associated larger punctures. Median portion of clypeus 

not raised nor projecting anteriorly as a lobe . . PLATYTHYREA Roger (p. 331) 

- Tibiae of middle and hind legs each with one large pectinate and one small simple 

spur. Sculpture not as above. Median portion of clypeus abruptly raised and 
projecting forwards as a truncated lobe . . . PALTOTHYREUS Mayr (p. 338) 

18 Middle and hind tibiae each with a single pectinate spur ..... 19 

- Middle and hind tibiae each with two spurs, one large and pectinate, the other small 

and simple ............. 21 

19 Basal portion of mandible with a distinct pit or fovea dorsolaterally 

CRYPTOPONE Emery (p. 336) 

- Basal portion of mandible without a dorsolateral pit or fovea .... 20 

20 Larger ants, total length greater than 7 mm. Mandibles elongate-falcate, their 

apical margins edentate or crenulate, the apex prolonged so that the tips cross 

over at rest PSALIDOMYRMEX E. Andre (p. 339) 

- Smaller ants, total length less than 5 mm. Mandibles triangular with 3-4 enlarged 

teeth apically, usually followed by a row of denticles. Apical margins meet 
evenly when mandibles at rest . . . HYPOPONERA Santschi (p. 336) 

21 Petiole thick, nodiform. Metanotal suture obsolete or absent, not impressed. 

Propodeum not or only slightly narrower than pronotum in dorsal view 

BOTHROPONERA Mayr (p. 334) 

- Petiole a thick scale. Metanotal suture present, usually distinct and impressed. 

Propodeum narrower than pronotum in dorsal view ..... 22 

22 Basal portion of mandible with a dorsolateral pit or fovea ..... 23 

- Basal portion of mandible without a dorsolateral pit or fovea 

MESOPONERA Emery (p. 338) 

23 Palp formula 3,3. Eyes larger, their maximum diameter greater than the greatest 

width of the scape. Metanotal suture very deeply impressed 

BRACHYPONERA Emery (p. 335) 

- Palp formula 4,4. Eyes smaller, their maximum diameter less than the greatest 

width of the scape. Metanotal suture very weakly impressed 

TRACHYMESOPUS Emery (p. 340) 



ANT GENERA OF WEST AFRICA 



325 



Subfamily CERAPACHYINAE 

1 Eyes absent. Gastral segments separated from each other by distinct constrictions 

SPHINCTOMYRMEX Mayr (p. 341) 

- Eyes present, varying from large to minute. Gaster constricted only between first 

and second segments, sometimes very deeply so, so that the pedicel is to all intents 

and purposes two-segmented .......... 2 

2 Tibial spurs absent from middle legs. Claws toothed. Antennal scapes much 

compressed dorsoventrally, more or less triangular in dorsal view 

SIMOPONE Forel (p. 341) 

3 Tibial spurs present on middle legs. Claws simple. Antennal scapes not or only 

slightly compressed, not triangular in dorsal view ...... 3 

- Apical segment of funiculus strongly swollen, forming a one-jointed club. Petiole 

not marginate laterally, never armed with teeth posterodorsally 

CERAPACHYS F. Smith (p. 340) 

- Apical three funicular segments swollen, forming a weak club. Petiole sharply 

marginate laterally, often with a pair of teeth situated posterodorsally 

PHYRACACES Emery (p. 340) 



MYRMICHRIA Saunders (p. 



Subfamily MYRMICINAE 

Antennae with 4 segments 

Antennae with 6 segments 

Antennae with 7 segments 

Antennae with 9 segments 

Antennae with 10 segments 

Antennae with 1 1 segments 

Antennae with 12 segments 

Mandibles short, subtriangular, serially dentate. Clypeus projecting anteriorly and 

overhanging mandibles for most of their length. Head without specialized 

orbicular hairs MICCOSTRUMA Brown (p. 

Mandibles elongate, linear, not serially dentate. Clypeus not projecting anteriorly 

over the mandibles. Head with numerous specialized orbicular hairs 
Apex of each mandibular blade with a fork of two spiniform teeth. Labrum not 

produced between mandibles as a biconical structure 

QUADRISTRUMA Brown (p. 
Apex of each mandible with a vertical series of denticulae. Labrum produced 

between mandibles as a biconical structure . . . EPITRITUS Emery (p. 

Mandibles long and linear, with not more than 4 teeth on the inner margin of each 

blade .............. 

Mandibles short, triangular or subtriangular, serially dentate or denticulate . 
Mandibles with an apical fork of 2-3 teeth, which may have one or more intercalary 

denticulae. Head without orbicular hairs ....... 

Mandibles without apical fork; either with a single long tooth at the dorsal apex 

subtended by a series of denticulae or with a series of denticulae only. Head 

with orbicular hairs present ..... EPITRITUS Emery (p. 

Antennal scrobes absent. Mandibular blades without preapical teeth. Occipital 

lobes of head, promesonotum, propodeum and petiole armed with teeth or short 

spines MICRODACETON Santschi (p. 

Antennal scrobes present above the eyes. Mandibular blades with one or two 

preapical teeth or denticles. Propodeum only armed with spines 

STRUMIGENYS F. Smith (p. 
Antennal scrobes absent. Propodeum unarmed, smoothly rounded. First (basal) 

tarsal segment on each leg swollen, as broad as tibiae 

MELISSOTARSUS Emery (p. 



2 

4 

349) 

11 

M 
17 
26 



343) 
3 

344) 
343) 

5 

7 

6 
343) 
344) 
345) 
347) 



326 B. BOLTON 

- Antenna] scrobes present above the eyes. Propodeum armed with a pair of spines 

or teeth. First tarsal segment on each leg slender ..... 8 

8 Apical border of mandible with more than 20 denticulae, the basal 4 to 8 of which 

may be considerably larger than those preceding. SERRASTRUMA Brown (p. 344) 

- Apical border of mandible with less than 10 teeth of variable size ... 9 

9 Head elongate, always with numerous setae . . . . . . . 10 

- Head not elongate, setae reduced to 2 or 3 on posterior dorsum 

TRICHOSCAPA Emery (p. 345) 

10 Clypeal hairs bizarre, either flattened, clavate or spoon-shaped 

SMITHISTRUMA Brown (p. 345) 

- Clypeal hairs not bizarre, usually short and simple. CODIOMYRMEX Wheeler (p. 343) 

11 Antennal scrobes present above the eyes; antennal club of three segments. Pro- 

mesonotum produced into a shelf posteriorly, dentate or spinose behind and over- 
hanging the propodeum, which is vertical or nearly so. Entire body densely 
clothed with fine hairs MERANOPLUS F. Smith (p. 348) 

- Antennal scrobes absent; antennal club of two segments. Promesonotum not 

produced posteriorly, propodeum not vertical. Hairs sparse . . . . 12 

12 Propodeum armed with a pair of spines or teeth or sharply angled. Worker caste 

dimorphic, without intermediate forms . . . OLIGOMYRMEX Mayr (p. 352) 

- Propodeum unarmed. Worker caste monomorphic ...... 13 

13 Eyes completely absent; mandibles armed with 5-6 teeth. Promesonotum not 

marginate laterally ...... CAREBARA Westwood (p. 351) 

- Eyes present, small, of 2 ommatidia only; mandibles with 4 teeth. Promesonotum 

distinctly marginate laterally .... PAEDALGUS Forel (p. 353) 

14 Petiole without a node. Postpetiole attached to dorsum of first gastral segment, 

the gaster itself more or less heart-shaped in dorsal view and capable of reflexion 

over the alitrunk CREMATOGASTER Lund (p. 342) 

- Petiole with a node. Postpetiole not attached to dorsum of first gastral segment; 

gaster not heart-shaped nor capable of reflexion over alitrunk . . . . 15 

15 Antennal scrobes present above the eyes; antennal club 3-segmented. Clypeus not 

longitudinally bicarinate. Sting with a triangular lamella projecting apico- 
dorsally DECAMORIUM Forel (p. 355) 

- Antennal scrobes absent; antennal club 2-segmented. Clypeus longitudinally 

bicarinate. Sting without triangular lamelliform appendage . . . . 16 

16 Propodeum smoothly rounded, neither sharply angulate nor dentate. Maxillary 

palpi geniculate. Mono- or polymorphic . SOLENOPSIS Westwood (p. 354) 

- Propodeum sharply angled or armed with a pair of spines or teeth. Maxillary 

palpi not geniculate. Dimorphic without intermediates. 

OLIGOMYRMEX Mayr (p. 352) 

17 Postpetiole attached to dorsum of first gastral segment. Gaster more or less heart- 

shaped and capable of reflexion over the alitrunk. 

CREMATOGASTER Lund (p. 342) 

- Postpetiole not attached to dorsum of first gastral segment. Gaster not heart- 

shaped nor capable of reflexion over the alitrunk . . . . . .18 

18 Antennal scrobes present, either above or below the eyes . . . . . 19 

- Antennal scrobes absent ........... 23 

19 Antennal scrobes below the eyes. Head, alitrunk and gaster dorsoventrally 

flattened, and often armed with spines or teeth along the sides of the head and 
alitrunk. First tergite forming the whole dorsal surface of the gaster when 
viewed from above CATAVLACUS F. Smith (p. 342) 

- Antennal scrobes above eyes. Head, alitrunk and gaster not dorsoventrally 

flattened. More than one gastral tergite visible in dorsal view. ... 20 

20 Antennal scrobes weak, bordered laterally by a longitudinal genal carina. Minute 

ants, 2 mm or less in total length .... WASMANNIA F. Smith (p. 350) 



ANT GENERA OF WEST AFRICA 327 

- Antennal scrobes more definitely marked, not bordered by lateral genal carinae. 

Larger, total length more than 2 mm . . . . . . . .21 

21 Median portion of clypeus vertical, with a distinct anteriorly projecting bilobed 

appendage dorsally; lateral portions of clypeus entire, not reduced to a narrow 
strip nor raised into a ridge in front of the antennal insertions. Head and body 
usually with some spatulate or otherwise bizarre hairs 

CALYPTOMYRMEX Emery (p. 347) 

- Median portion of clypeus not vertical, without a bilobed appendage dorsally. 

Lateral portions of clypeus in front of antennal insertions reduced to a narrow 
strip or raised into a ridge. Head and body without spatulate or otherwise 
bizarre hairs ............ 22 

22 Sting with a spatulate lamella apically. Anterior margin of median portion of 

clypeus without projections. Pronotum always unarmed; petiole sometimes 
squamiform XIPHOMYRMEX Forel (p. 356) 

- Sting acute, not ending in a spatulate lamella apically. Anterior margin of median 

portion of clypeus with a number of low, blunt, dentiform projections. Pronotum 
armed above with a pair of spines or a pair of low blunt tubercles ; petiole never 
squamiform PRISTOMYRMEX Mayr (p. 348) 

23 Antenna] club of 2 segments. Propodeum armed with a pair of spines or teeth . 24 

- Antennal club of 3 segments. Propodeum smoothly rounded, unarmed . . 25 

24 Clypeus bicarinate. Dimorphic, without intermediates 

OLIGOMYRMEX Mayr (p. 352) 

- Clypeus not bicarinate. Polymorphic, with a graded series of intermediates 

connecting minor to major workers ' . . PHEIDOLOGETON Mayr (p. 353) 

25 Median portion of clypeus distinctly longitudinally bicarinate 

MONOMOR1UM M&yr (p. 351) 

- Median portion of clypeus somewhat swollen but not bicarinate 

DIPLOMORIUM Mayr (p. 351) 

26 Petiole armed with a pair of spines or teeth above ...... 27 

- Petiole unarmed or merely emarginate above ....... 28 

27 Mesonotum bituberculate; propodeum armed with a pair of long spines 

ATOPOMYRMEX E. Andre (p. 348) 

- Mesonotum not bituberculate; propodeum unarmed . TERATANER Emery (p. 349) 

28 Spurs on middle and hind tibiae pectinate . . CRATOMYRMEX Emery (p. 349) 

- Spurs on middle and hind tibiae simple or absent ...... 29 

29 Antennal club of 2 segments, the apical segment greatly enlarged. Clypeus 

produced into a lobe anteriorly. Basal internal margin of each mandible with a 
tooth ADELOMYRMEX Emery (p. 346) 

- Antennal club of 3 or 4 segments, or absent. Basal internal margin of each mandible 

without a tooth ............ 30 

30 Lateral portions of clypeus with their posterior margins raised into a ridge in front 

of the antennal insertions. Sting with a lamelliform triangular appendage 
apicodorsally. 

Propodeum usually bispinose and with a pair of metapleural lobes or teeth 
below the propodeal spines . . . . . . . . . .31 

- Lateral portions of clypeus not raised into a ridge in front of the antennal insertions. 

Sting without a lamelliform appendage apicodorsally ..... 33 

31 Head and body with trifid or multifid setae, usually abundant. (In most cases the 

trifid setae are best observed at mag. x 80 or above by viewing the alitrunk and 
pedicel in profile.) Antennal scrobes usually present, deep, divided into upper 
and lower compartments by a median longitudinal ridge which is incomplete 
posteriorly TRIGLYPHOTHRIX Forel (p. 356) 

- Head and body usually with scattered simple setae, very rarely with multifid setae 

or without setae. Scrobes present or absent; when present, never as above . 32 



328 B. BOLTON 

32 Propodeum armed with a pair of spines or teeth of varying configuration. Meta- 

pleural lobes or teeth present. Palp formula usually 4,3. 

TETRAMORIUM Mayr (p. 355) 

- Propodeum unarmed, rounded. Metapleural lobes present. Palp formula 3,2. 

RHOPTROMYRMEX Mayr (p. 355) 

33 Antennal scrobes present above the eyes. Median portion of clypeus vertical, with 

a dorsal bilobed appendage projecting anteriorly. Usually with bizarre setae 
present on head, alitrunk and gaster . . CALYPTOMYRMEX Emery (p. 347) 

- Antennal scrobes absent. Median portion of clypeus not vertical, without a bilobed 

appendage. Bizarre setae absent ......... 34 

34 Propodeum unarmed, smoothly rounded ........ 35 

- Propodeum armed with a pair of spines or teeth ...... 36 

35 Clypeus with median portion distinctly longitudinally bicarinate. 

MONOMORIUM Mayr (p. 351) 

- Clypeus with median portion rounded, not bicarinate. 

SYLLOPHOPSIS Santschi (p. 355) 

36 Gular surface of head with a psammophore. Polymorphic . MESSOR Forel (p. 350) 

- Gular surface of head without a psammophore. Monomorphic, or dimorphic 

without intermediates . . . . . . . . . . -37 

37 Anterodorsal angles of pronotum sharply angled in dorsal view. Body hairs either 

clavate or short and very stout. Petiole with a very short, broad peduncle 
anteriorly LEPTO THORAX Mayr (p. 346) 

- Anterodorsal angles of pronotum rounded in dorsal view. Body hairs either sparse 

or abundant and long, but always simple. Petiole with a long anterior peduncle 38 

38 Dimorphic, without intermediates. In both castes petiolar node antero-posteriorly 

compressed, conical or subconical in profile and with the dorsal border usually 
emarginate. Major workers with head emarginate behind and with crushing 
mandibles usually equipped with 3 teeth, of which two are situated apically and 
one basally, separated by a large diastema. In the minor worker the sides of the 
head usually converge behind the eyes so that the occipital margin is short 

PHEIDOLE Westwood (p. 350) 

- Monomorphic. Petiolar node in profile domed or flattened above, never conical or 

subconical in shape, never emarginate above ....... 39 

39 Antennal scapes short, not surpassing occipital margin of head. Lateral portions 

of clypeus projecting forwards over the base of the mandibles. Pronotum 
smoothly rounded, not tuberculate. Alitrunk and gaster devoid of long setae 

CARDIOCONDYLA Emery (p. 341) 

- Antennal scapes very long, easily surpassing occipital margin of head. Lateral 

portions of clypeus not projecting forwards over base of mandibles. Pronotum 
with a small obtuse tubercle on each side. Entire body clothed with long erect 
setae MACROMISCHOIDES Wheeler (p. 347) 

Subfamily DORYLINAE 

1 Pedicel of 2 segments. Antennae 10-segmented; genae carinate. Pygidium not 

impressed, not armed with spines .... AENICTUS Shuckard (p. 357) 

- Pedicel of a single segment. Antennae 9- to 11-segmented: genae not carinate. 

Pygidium impressed, armed with a short spine or tooth at each side posteriorly. 

DORYLUS Fabricius (p. 357) 

Subfamily LEPTANILLINAE 

- With a single genus in West Africa, answering the characters given in the key to 

subfamilies LEP TA NILL A Emery (p. 358) 



ANT GENERA OF WEST AFRICA 329 

Subfamily PSEUDOMYRMECINAE 

Claws toothed. Palp formula 5,4. Funiculi of antennae not clubbed apically. 

Eyes occupying one third or more of the side of the head .... 2 

Claws simple. Palp formula 3,3. Funiculi with a club of three segments. Eyes 

smaller, occupying approximately one fifth of the side of the head 

VITICICOLA Wheeler (p. 359) 
Both petiole and postpetiole with strong ventral processes. Eyes occupying about 

one third the side of the head. 3 ocelli present . PACHYS1MA Emery (p. 358) 
Petiole only with a weak ventral process, or both nodes without processes. Eyes 

occupying one third to one half the side of the head. Ocelli variable, often 

absent, but may be one, two or three developed TETRAPONERA F. Smith (p. 358) 

Subfamily DOLICHODERINAE 

Scale of petiole well developed, distinct, inclined anteriorly but not reduced nor 

overhung by first gastral segment dorsally . JRIDOMYRMEX Mayr (p. 359) 

Scale of petiole reduced or vestigial; petiole overhung by first gastral segment 

dorsally, usually invisible in dorsal view ....... 2 

Palp formula 4,3, palpi short ...... ENGRAMMA Forel (p. 359) 

Palp formula 6,4, palpi long .......... 3 

In dorsal view 5 gastral segments visible. Anal and associated orifices apical 

TECHNOMYRMEX Mayr (p. 360) 
In dorsal view 4 gastral segments visible. Anal and associated orifices ventral 

TAPINOMA Forster (p. 359) 

Subfamily FORMICINAE 

Antennae 9-segmented ..... APHOMOMYRMEX Emery (p. 362) 

Antennae 11 -segmented ........... 2 

Antennae 12-segmented ........... 4 

Palp formula 1,3 or 2,3 Eyes minute. 

Small hypogaeic ants ...... ACROPYGA Roger (p. 363) 

Palp formula 6,4. Eyes distinct; ocelli may be present ..... 3 

Propodeum bidentate or bituberculate; petiole usually bispinose or bidentate above, 

occasionally only with upper border strongly emarginate 

ACANTHOLEPIS Mayr (p. 363) 
Propodeum unarmed; petiole neither armed nor emarginate above. 

PLAGIOLEPIS Mayr (p. 364) 
Eyes enormous, occupying almost all the side of the head. Ventrolateral margin 

of head with a tooth at each side. Propodeum bispinose 

SANTSCHIELLA Forel (p. 364) 
Eyes smaller, occupying less than one half the length of the side of the head. 

Ventrolateral margin of head unarmed ; propodeum usually unarmed . . 5 

Ocelli present. Psammophore present on the anterior portion of the gular surface 

of the head CAT A GLYPH IS Forster (p. 361) 

Ocelli vestigial or absent; psammophore absent ....... 6 

Petiole reduced to an elongate, low node, allowing the gaster to be reflexed over the 

alitrunk. Mandibles elongate triangular, broad, the apical tooth long. Palp 

formula 5,4 OECOPHYLLA F. Smith (p. 363) 

Petiole a node or scale, never as above; mandibles not as above. Palp formula 

usually 6,4, rarely reduced to 3,4 or 3,3, but never 5,4 .... 7 

Antennal insertions very close to, or contiguous with the posterior clypeal margin. 

Acidopore borne on a conical projection of the hypopygium, forming a nozzle, the 

orifice surrounded by a fringe of hairs ........ 8 



33o B. BOLTON 

— Antennal insertions some distance (usually greater than basal width of scape) 

behind the posterior clypeal margin. Acidopore not borne on a conical projection 
of the hypopygium, the orifice usually not surrounded by a fringe of hairs; or the 
acidopore hidden by a projection of the pygidium ...... 10 

8 Polymorphic. Eyes small in major workers, absent in minors. Clypeus over- 

hanging mandibles in front. Palpi short, indistinct, palp formula reduced to 

3,4 in major and 3,3 in minor workers . . PSEUDOLASIUS Emery (p. 362) 

— Monomorphic. Eyes well developed, occasionally large. Clypeus not or only 

slightly overhanging the mandibles. Palp formula 6,4, the palpi long and distinct 9 

9 Dorsum of alitrunk with very coarse setae arranged in distinct pairs. Eyes at or in 

front of the midlength of the head . PARATRECHINA Motschulsky (p. 361) 

— Dorsum of alitrunk with fine setae, not definitely paired. Eyes behind the mid- 

length of the head PRENOLEPIS Mayr (p. 362) 

10 Anterodorsal pronotal angles usually projecting as spines or teeth, at least strongly 

marginate. Propodeum usually bispinose or bidentate; petiole with sharp 
angles, spines or teeth above. Monomorphic . . . . . . . 11 

— Anterodorsal pronotal angles rounded, unarmed. Propodeum unarmed although 

may be truncated posteriorly. Petiole a node or scale, never with teeth or spines. 
Polymorphic CAMPONOTUS Mayr (p. 360) 

11 Clypeus flat, not longitudinally carinate in the middle, its anterior margin broadly 

and shallowly emarginate. Acidopore not concealed by the pygidium 

PHASMOMYRMEX Stitz (p. 360) 

— Clypeus usually convex and longitudinally carinate, its anterior margin usually not 

emarginate. Acidopore concealed by the pygidium when not in use 

POLYRHACHIS F. Smith (p. 361) 

SYNONYMIC SYNOPSIS AND NOTES ON THE GENERA 

Subfamily PONERINAE 
Tribe AMBLYOPONINI 
AMBLYOPONE Erichson 

Amblyopone Erichson, 1842, Arch. Naturgesch. 8(1) : 260. Type-species: Amblyopone australis 

Erichson, 1842, op. cit. : 261, by monotypy. 
Stigmatomma Roger, 1859, Berl. ent. Z. 3 : 250. Type-species: Stigmatomma denticulatum 

Roger, 1859, op. cit. : 251, by designation of Bingham, 1903. 
Arotropus Provancher, 1881, Can. Nat. 6- Geol. 12 : 205. Type-species: Arotropus binodosus 

Provancher, 1881, by monotypy. ( = Amblyopone pallipes (Haldeman) 1844, Proc. Acad. 

nat. Sci. Philad. 2 : 54.) 
Xymmer Santschi, 1914, Boll. Lab. Zool. gen. agr. Portici 8 : 311 [as a subgenus of Stigmatomma]. 

Type-species: Stigmatomma (Xymmer) muticum Santschi, 1914, loc. cit., by monotypy. 
Fulakora Mann, 1919, Bull. Mus. comp. Zool. Harv. 63 : 279 [as a subgenus of Stigmatomma]. 

Type-species: Stigmatomma (Fulakora) celata Mann, 1919, loc. cit., by original designation. 
Neoamblyopone Clark, 1927, in Wheeler, Proc. Am. Acad. Arts Sci. 62 : 1 [as a subgenus of 

Amblyopone]. Type-species: Amblyopone (Neoamblyopone) clarki Wheeler, 1927, op. cit. : 

24, by monotypy. 
Protamblyopone Clark, 1927, in Wheeler, Proc. Am. Acad. Arts Sci. 62 : 1 [as a subgenus of 

Amblyopone]. Type-species: Amblyopone (Protamblyopone) aberrans Wheeler, 1927, op. 

cit. : 26, by monotypy. 
Lithomyrmex Clark, 1928, /. Proc. R. Soc. West. Aust. 14 : 30-31. Type-species: Lithomyrmex 

glauerti Clark, 1928, loc. cit., by original designation. 
Ericapelta Kusnezov, 1955, Zool. Anz. 154 : 273-274. Type-species: Ericapelta egregia 

Kusnezov, 1955, loc. cit., by monotypy. 



ANT GENERA OF WEST AFRICA 331 

The mandibles are long and narrow, inserted far apart at the corners of the clypeus and 
tending to cross over at rest ; their apices acute and their inner margins dentate throughout the 
length of the blades. The median portion of the clypeus projects anteriorly as a lobe which 
usually bears a series of denticulae, but in A . mutica (Santschi) the lobe is straight and edentate 
in front. An anterolateral dentiform projection of the gena may be present. On the alitrunk 
the promesonotal suture is complete, the metanotal may be present or absent. Petiole massive, 
very broadly attached to the first gastral segment and without a free posterior face. Sculpture 
is usually fine; the head with fine dense puncturation which may be continued onto the pro- 
notum, or with the pronotum more or less smooth and shining. Palp formulae of 5,3 ; 4,3 ; 3,2 
and 2,2 have been recorded in the genus (Brown, i960). Workers are found in moist soil 
and have been recovered from leaf -litter samples by Berlese funnel separation. 

Amblyopone is represented in West Africa by a few rare and little known species, 
mostly with a savannah distribution. 

MYSTRIUM Roger 

Mystrium Roger, 1862, Berl. ent. Z. 6 : 245. Type-species: Mystrium mysticum Roger, 1862, 
loc. cit., by monotypy. 

Mandibles very long, noticeably longer than the head, broad and blunt apically, their inner 
margins with a double row of short teeth. The anterior margin of the median portion of the 
clypeus is denticulate, the genal teeth are well developed and project anteriorly. Structure of 
alitrunk and petiole as in Amblyopone but the metanotal suture represented by a weak impression 
dorsally, which does not break the sculpture. The tarsal segments possess numerous spiniform 
setae, most abundant on the internal surface of the basal segment, more evenly distributed 
on the remaining segments. Sculpture of a coarse reticulation over the head and alitrunk; 
all dorsal surfaces of head and body with spatulate setae present. Menozzi (1929) and Brown 
(i960) give a palp formula of 4,3 for the genus. 

Known only from a single species in West Africa, distributed throughout the 
forest zone, M. silvestrii Santschi has been noted in Ghana nesting in soil at the base 
of a cocoa tree in an old (abandoned) cocoa plantation. 

Tribe PLATYTHYREINI 

PLATYTHYREA Roger 

Platythyrea Roger, 1863, Berl. ent. Z. 7 : 172. Type-species : Pachycondyla punctata F. Smith, 
1858, Cat. Hym. Brit. Mus. 6 : 108, by designation of Bingham, 1903. 

This genus and the one below constitute the tribe Platythyreini as represented in 
West Africa, and are easily recognizable by the distinctive sculpture which both 
possess. 

Entirety of head, alitrunk, pedicel, gaster and appendages with very fine shagreening and 
with scattered larger, shallow punctures. All surfaces covered by an extremely fine dense 
pubescence and devoid of standing hairs. 

Platythyrea is further distinguished by the possession of two pectinate spurs on each of the 
middle and hind tibiae, and the presence of a median tooth on the pretarsal claws. Palp 
formulae of 6,4 and 3,2 have been recorded in West African species, the majority of which 
nest in rotten wood on the ground or in rotten parts of standing trees. The lobes of the frontal 
carinae are usually broad and fused to the clypeus in front. Clypeus itself is broad and convex, 
not carinate; eyes are always present although they may be reduced. Antennae 12-segmented, 



332 B. BOLTON 

often with the second funicular segment longer than the first or third. Alitrunk with prome- 
sonotal suture distinct, the metanotal obsolete or absent. Petiole a large node, often with the 
posterodorsal margin extended into dentiform projections. 

PROBOLOMYRMEX Mayr 

Probolomyrmex Mayr, 1901, Annln naturh. Mus. Wien 16 : 2. Type-species: Probolomyrmex 

filiformis Mayr, 1901, op. cit. : 3, by monotypy. 
Escherichia Forel, 1910, Zool. Jb., Abt. Syst. 29 : 245. Type-species: Escherichia brevirostris 

Forel, 1910, op. cit. : 246, by monotypy. 

Sculpture is as in Platythyrea above; Probolomyrex has the following distinguishing features. 

Frontal carinae fused together and fused to the clypeus, projecting as a plate over the man- 
dibles. Antennae inserted close together and close to the anterior margin of this plate, their 
condylar bulbs exposed in dorsal view. Antennae 12-segmented, the funicular segments be- 
coming thicker towards the apex. Palp formula 4,2 (Taylor, 1965). Alitrunk without dorsal 
sutures. Middle and hind tibiae each with a single pectinate spur; claws simple. 

Probolomyrmex, a rare genus of small ants with a tropicopolitan distribution is 
represented in Africa by three species, one of which, P. guineensis Taylor occurs in 
West Africa. Nests are made in small pieces of rotted wood in cultivated or other- 
wise disturbed ground on the coastal plain. 

Tribe ECTATOMMINI 
DISCOTHYREA Roger 

Discothyrea Roger, 1863, Berl. ent. Z. 7 : 176. Type-species: Discothyrea testacea Roger, 

1863, op. cit. : 177, by monotypy. 
Pseudosysphincta Arnold, 1916, Ann. S. Afr. Mus. 14 (2) : 161. Type-species: Pseudosysphincta 

poweri Arnold, 1916, op. cit. : 162, by original designation. 
Prodiscothyrea Wheeler, 1916, Trans. R.Soc. West. Aust. 40 : 33. Type-species: Prodiscothyrea 

velutina Wheeler, 1916, op. cit. : 34, by monotypy. 
Pseudosphincta Wheeler, 1922, Bull. Am. Mus. nat. Hist. 45 : 645, 762. [Variant spelling of 

Pseudosysphincta.'] 

Characters shared by the two genera Discothyrea and Proceratium include reduction in size 
of the eyes and depigmentation of the body in response to cryptobiotic habits, raised frontal 
carinae leaving the condylar bulbs of the antennae exposed in dorsal view, lack of sutures on 
the dorsum of the alitrunk, and strong vaulting of the second gastral segment so that the remain- 
ing segments are directed anteriorly below the first and second. 

In Discothyrea the mandibles are overhung by the clypeus, and are edentate; the antennae 
are nine or eleven-segmented in known West African species, with the apical funicular segment 
strongly swollen and bulbous. Spurs on the middle tibiae small and simple, those on the 
hind tibiae pectinate. Sculpture of dense puncturation over most of the body. 

Both species found in West Africa, D. oculata Emery with nine-segmented anten- 
nae, and D. mixta Brown with eleven-segmented antennae, are apparently confined 
to the forest zone. D. oculata is found throughout the region although quite rarely, 
whilst D. mixta was originally described from Liberia and is now also known to 
occur in Ghana. 

Discothyrea and the closely related genus Proceratium represent the tribe Ectatom- 
mini in West Africa. The former genus has a southerly distribution, whilst the 



ANT GENERA OF WEST AFRICA 333 

latter is mostly confined to the northern hemisphere, but in the tropics their ranges 
tend to overlap. Both genera are cryptobiotic and form small colonies, usually 
in rotten wood just below the surface of the ground. Workers have also been 
recovered from Berlese funnel samples of leaf-litter. 

PROCERATIUM Roger 

Proceratium Roger, 1863, Berl. ent. Z. 7 : 171. Type-species: Proceratium silaceum Roger, 

1863, op. cit. : 172, by monotypy. 
Sysphingta Roger, 1863, Berl. ent. Z. 7 : 175. Type-species: Sysphingta micrommata Roger, 

1863, op. cit. : 176, by monotypy. 
Sysphincta Mayr, 1865, Reise der . . . Fregatte Novara, Zool. 2 : 12. Wien. [Emendation of 
Sysphingta.'] 

Known from West Africa by a single new species discovered at Legon, Ghana, in 
a small piece of rotten wood by D. Leston, and to be described elsewhere by him. 

Proceratium shares a number of characters with Discothyrea as mentioned above but has 
the mandibles armed with three teeth, only slightly overhung by the clypeus, the antennae 
12-segmented with the apical segment enlarged but not bulbous. Brown (1958) records palp 
formulae of 4,3 ; 3,2 ; and 2,2 in the genus; that of the West African species is found to be 4,3. 

Tribe ODONTOMACHINI 
ODONTOMACHUS Latreille 

Odontomachus Latreille, 1805, Hist. Nat. Crast. Ins. 13 : 257. Type-species: Formica haematoda 

Linnaeus, 1758, Syst. Nat. ed. 10 : 582, by original designation. 
Pedetes Bernstein, 1861, Verh. zool.-bot. Ges. Wien 11 : 7. Type-species: Pedetes macrorhynchus 

Bernstein, 1861. [Nomen nudum.] 

Mandibles elongate, linear, inserted close to the midline of the anterior margin of the head 
and meeting throughout their length when at rest. When fully open the mandibles form a 
line at right angles to the long axis of the head. Apical mandibular armament of three teeth 
arranged in a vertical series, the dorsalmost tooth truncated. Eyes situated close to the 
anterior margin of the head, separated from the frontal carinae by a broad longitudinal impres- 
sion. Palp formula 4,4 or 4,3. Node of petiole ending in a long spine dorsally; gaster not 
impressed between first and second segments. 

Of the two species found in West Africa, 0. haematodus (L.) prefers open or loosely 
wooded areas whilst 0. assiniensis Emery is more commonly found in thick bush 
and more dense forest. Both species nest in rotten wood at ground level, and 
0. haematodus may nest at the bases of trees, especially in cultivated areas. When 
the nests are in such a location the ants often ascend the tree to forage, one of the 
few ponerine species to do so in West Africa. 

A NO CHE TUS Mayr 

Anochetus Mayr, 1861, Europ. Formicid. : 53. Type-species: Odontomachus ghilianii Spinola, 
1851, Memorie Accad. Sci. Torino (2)13 : 71, by designation of Bingham, 1903. 

Myrmapatetes Wheeler, 1929, Am. Mus. Novit. no. 349 : 6. Type-species: Myrmapatetes 
filicornis Wheeler, 1929, loc. cit., by original designation. 



334 B. BOLTON 

Answering to the description given above for Odontomachus but the apical mandibular 
armament with the dorsalmost tooth acute, the palp formula 4,3 and the petiolar node not 
ending in a spine dorsally. Whilst these characters hold good for the species of West Africa 
they do not, unfortunately, apply throughout the world and Anochetus is at best a very weak 
genus, hardly if at all separable from Odontomachus. 

The majority of species ate found in leaf-litter and rotten wood, but one species 
is known to nest and forage arborially. 

Tribe PONERINI 
ASPHINCTOPONE Santschi 

Asphinctopone Santschi, 1914, Boll. Lab. Zool. gen. agr. Portici 8 : 318. Type-species: Asphinc- 

topone silvestrii Santschi, 1914, loc. cit., by monotypy. 
Lepidopone Bernard, 1952, Mem. Inst, f rang. Afr. noire no. 19 : 207. Type-species: Lepidopone 

lamottei Bernard, 1952, op. cit. : 208, by monotypy. 

Mandibles somewhat elongate, armed with five teeth, the basal internal margin with a 
distinct notch just before the articulation, which is concealed by the clypeus when the mandibles 
are completely closed. Clypeus bluntly carinate, the median portion projecting as a rectangular 
lobe. Eyes reduced. Dorsum of alitrunk with promesonotal suture and metanotal groove 
present, distinct, the latter very deeply impressed, with longitudinal ribbing. Declivity of 
propodeum steep, marginate at sides and above. Petiole a high, narrow scale, the posterior 
peduncle with a number of distinct raised transverse ridges. Subpetiolar process complex, 
with three prominences. Gaster with a large an tero ventral process on the first segment; not 
constricted between the first and second segments. 

A . silvestrii Santschi has the head finely and densely punctate, the pronotum sparsely punctate 
with wide shining interspaces and a virtual lack of puncturation on the mesonotum. Propodeum 
with sparse, more coarse punctures, considerably larger than those on the pronotum. 

This rarely found genus is closely related to Mesoponera. The specimens before 
me were found in a black-rotten, very wet banana trunk, near Ibadan, Western 
Nigeria. 

BOTHROPONERA Mayr 

Bothroponera Mayr, 1862, Verh. zool.-bot. Ges. Wien. 12 : 717. Type-species: Ponera pumicosa 
Roger, i860, Berl. ent. Z. 4 : 29, by designation of Emery, 1901, Annls Soc. ent. Belg. 45 : 45. 

Pseudoneoponera Donisthorpe, 1943, Ann. Mag. nat. Hist. (n)10 : 439. Type-species: 
Pseudoneoponera verecundae Donisthorpe, 1943, loc. cit., by original designation. 

Medium to large ants, usually quite coarsely sculptured and black in colour, occasionally 
dark brown or deep red-brown. Mandibles dentate, usually with six or seven teeth, reduced to 
four or five in the B. nasica Santschi group. B. sjoestedti (Mayr) has a mandibular fovea 
situated basally on the dorsolateral surface as in Brachyponera, Trachymesopus and Cryptopone. 
Palp formula usually 4,4, reduced to 2,2 in B. sjoestedti and the B. nasica group. Alitrunk 
with promesonotal suture present, mobile, the metanotal absent. In dorsal view the propo- 
deum is scarcely or not narrower than the pronotum. Petiole thick and nodiform. Middle 
and hind tibiae each with a large pectinate and a small simple spur, the latter may be very 
much reduced in members of the B. nasica species-group. 

The majority of Bothroponera species are bush and forest inhabiting ants, nesting 
in rotten wood or directly into hard-packed earth. The larger species are mostly 



ANT GENERA OF WEST AFRICA 335 

free foragers and often nocturnal, but the smaller forms are more cryptic and are 
often found in association with termites. Most free foraging species prefer well 
shaded and moist habitats and are very rarely found in the open. 

BRACHYPONERA Emery 

Brachyponera Emery, 1901, Annls Soc. ent. Belg. 45 : 43 [as a subgenus of Euponera.} Type- 
species: Ponera senaarensis Mayr, 1862, Verh. zool.-bot. Ges. Wien. 12 : 721, by original 
designation. 

Brachyponera Emery; Wilson, 1958, Bull. Mus. comp. Zool. Harv. 119 : 346. [Raised to genus.] 

Represented by a single species B. senaarensis (Mayr) in West Africa. 

Mandibles with a distinct, oval pit or fovea on the dorsolateral surface. Palp formula 3,3. 
Eyes quite large, their maximum diameter distinctly greater than the maximum width of the 
scape. Promesonotal suture distinct, metanotal groove distinct and deeply impressed. Pro- 
podeum narrowed dorsally, in dorsal view narrower than the pronotum. Petiole a thick 
scale; gaster weakly impressed between first and second segments. Extremely finely and 
densely punctate everywhere. 

B. senaarensis is essentially a savannah species, but penetrates the forest zone 
bordering on savannah. Nests are made directly into the soil, usually in direct 
sunlight which the species makes no attempt to avoid. It is present throughout 
the West African savannah. 



CACOPONE Santschi 

Cacopone Santschi, 1914, Boll. Lab. Zool. gen. agr. Portici 8 : 325. Type-species : Cacopone 
hastifer Santschi, 1914, loc. cit., by monotypy. 

A monotypic genus very close to and probably inseparable from Plectrodena. 
It shares the characteristic mandibular form and articulation with Plectrodena 
(see below) and only differs from that genus in details of mandibular structure and 
nodal form. 

Mandibles elongate, linear, the internal margins edentate but with the distal half distinctly 
swollen and then narrowing to the apex. Eyes very much reduced and depigmented, difficult 
to see. Petiole node without a distinct anterior face, in profile this surface rounding convexly 
into the dorsal surface. 

C. hastifer Santschi is a little known species whose life-way is apparently com- 
pletely subterranean ; confined to forested areas. 



CENTROMYRMEX Mayr 

Centromyrmex Mayr, 1866, Verh. zool.-bot. Ges. Wien 16 : 894. Type-species: Centromyrmex 

bohemanni Mayr, 1866, op. cit. : 895, by monotypy. 
Spalacomyrmex Emery, 1889, Annali Mus. civ. Stor. nat. Giacoma Doria 27 : 489. Type-species: 

Spalacomyrmex feae Emery, 1889, loc. cit., by monotypy. 
Typholoteras Karawajew, 1925, Konowia 4 : 128. Type-species: Typhloteras hamulatum 

Karawajew, 1925, op. cit. : 129, by monotypy. 



336 B. BOLTON 

Glyphopone Forel, 1913, Revue Zool. afr. 2 : 308. Type-species: Glyphopone bequaerti Forel, 
1913, loc. cit., by monotypy. 

Leptopone Arnold, 1916, Ann. S. Afr. Mus. 14 : 163 [as a subgenus of Glyphopone']. Type- 
species: Glyphopone [Leptopone) rufigaster Arnold, 1916, loc. cit. [= Centromyrmex bequaerti 
(Forel, 1913)], by original designation. 



A single West African species, C. sellaris Mayr. 



Mandibles strongly downcurved; median portion of clypeus almost vertical; eyes absent. 
Pronotum and mesonotum flat dorsally, the former strongly margined anteriorly and laterally 
Metanotal groove absent but propodeum pinched in, and strongly concave at about the middle 
of its length in lateral view. Propodeum convex behind this impression and rounding into an 
almost straight and vertical declivitous face. Subpetiolar process a simple spine; gaster not 
constricted between first and second segments. Coxae large, those of the anterior legs relatively 
enormous. Tarsal segments on all legs equipped with numerous downcurved spines and stiff 
setae, present also on the extensor (outer) surfaces of the middle tibiae and the apices of the 
hind tibiae. Apical spurs of tibiae of middle legs both small and simple; hind tibiae with one 
large pectinate and one small simple spur. 

C. sellaris is a totally subterranean ant usually found with termites either in or 
under rotten logs which the termites are eating, or moving through galleries in the 
outer walls of the nests of mound-building termites Occasionally it is found in 
hard-packed earth above colonies of termites which do not raise mounds. The 
abundantly spiny tarsi of this species give traction on the walls of underground 
tunnels and are found (less well developed) in other, unrelated genera which have 
similar habits. 

CRYPTOPONE Emery 

Cryptopone Emery, 1892, Annls Soc. ent. Fr. 61 : 275. Type-species: Cryptopone testacea 
Emery, 1893, Annls Soc. ent. Fr. 62 : 240-241, nee ? Amblyopone testacea Motschulsky. 
1863, Bull. Soc. Nat. Moscou 36 : 15, by designation of Wilson, 1958, Bull. Mus. comp. Zool. 
Harv. 119 (4) : 360-361. 

Small ants (less than 4 mm), yellowish or ferruginous in colour. Apical mandibular margin 
armed with four to six teeth; basally the mandible with a distinct pit or fovea on the dorso- 
lateral surface. Middle tibiae with stout spinules on the extensor surface. Eyes minute or 
absent. Palp formula 2,2 or less (Brown, 1963). 

The two species originally described from Africa in this genus were removed to 
Ponera by Brown (1963 : 6) and later to Hypoponera by Taylor (1967 : 11), who 
also pointed out that a number of small African species of Hypoponera are convergent 
upon Cryptopone. Brown (1963 : 4) indicates that ponerine ants possessing a 
basal mandibular pit or fovea are chiefly of African origin and expresses surprise 
that 'no true Cryptopone are known to occur in Ethiopian Africa'. The genus is 
included in the present study on the assumption that specimens will eventually be 
discovered in sub-Saharan Africa. 

HYPOPONERA Santschi 

Hypoponera Santschi, 1938, Bull. Soc. ent. Fr. 43 : 78-80 [as a subgenus of Ponera]. Type- 
species: Ponera abeillei E. Andre, 1881, Annls Soc. ent. Fr. (6)1 : 48, by original designation. 
Hypoponera Santschi; Taylor, 1967, Pacif. Insects Monogr. 13 : 9-12. [Raised to genus.] 



ANT GENERA OF WEST AFRICA 337 

Small ants superficially similar to Cryptopone above. Mandibles armed with three or four 
teeth apically, usually followed by a series of denticulae, without a basal mandibular pit. Palp 
formula 1,1, or 1,2 (Taylor, 1967). Eyes reduced, usually present but absent in a few species. 
Middle and hind tibiae each with a single pectinate spur; no spinules present on extensor 
surfaces of middle tibiae. Sculpture usually of fine dense puncturation. 

Species of Hypoponera are common in West Africa, in leaf-litter and log-mould 
and are found nesting in fallen twigs, rotten logs (particularly when the bark is still 
present), compressed leaf-litter or hard-packed earth Colonies are small and the 
individual ants are relatively slow-moving. Some species look very similar to 
Cryptopone, as mentioned above, but the distinctions given are sufficient to separate 
the two genera. True Ponera, as defined by Taylor (1967: 5-9) does not occur in 
West Africa. 



LEPTOGENYS Roger 

Leptogenys Roger, 1861, Berl. ent. Z. 5 : 41. Type-species: Leptogenys falcigera Roger, 1861, 

op. cit. : 42, by designation of Bingham, 1903. 
Dorylozelus Forel, 1915, Ark. Zool. 9(ig) : 24-25. Type-species: Dorylozelusm joebevgi Forel, 

1915, loc. cit., by monotypy. 
Microbolbos Donisthorpe, 1948, Entomologist 81 : 170. Type-species: Microbolbos testaceus 

Donisthorpe, 1948, loc. cit., by original designation. 

Formerly included in its own tribe, Leptogenyini, which was synonymized to 
Ponerini by Brown (1963). 

Mandibles of varying shape, may be elongate, linear and curved, or short and quite broad but 
always more or less edentate, with only one or two teeth situated apically. Mandibles articul- 
ated at extreme corners of anterior margin of head. Median portion of clypeus carinate, 
produced anteriorly into a lobe or point. Palp formula 4,4. Lobes of frontal carinae small, 
usually only partially covering the condylar bulbs of the antennal scapes in dorsal view. Middle 
and hind tibiae each with one large pectinate and one small simple spur. Claws pectinate, 
incompletely so in some species. Gaster weakly impressed between first and second segments. 

The genus Leptogenys is distributed throughout West Africa but the greatest 
number of species inhabit the forest zone where they usually nest in wet-rotten 
wood or in hard-packed soil, and in leaf-litter under fallen tree trunks. 



MEGAPONERA Mayr 

Megaponera Mayr, 1862, Verh. zool.-bot. Ges. Wien 12 : 734. Type-species: Formica foetens 

Fabricius, 1793, Ent. Syst. 2 : 354, by monotypy. 
Megaloponera Roger, 1863, Verzeichniss Formiciden-Gattungen : 17. Type-species: Formica 

foetens Fabricius, 1793, Ent. Syst. 2 : 354, by monotypy. 

A monotypic genus, the species of which is confined to savannah regions in 
West Africa. 

Dimorphic. Palp formula 4,4. Clypeus with median area swollen but not carinate nor raised 
and projecting. Gena on each side with a distinct longitudinal carina running from the eye to 
the mandibular insertion. Middle and hind tibiae each with one large pectinate and one small 
simple spur; claws armed with a tooth near the base. 



338 B. BOLTON 

M. foetens (F.) has an aromatic odour and stridulates audibly when disturbed. 
Nests are made in the earth, from which columns of ants emerge to raid nearby 
termite nests. 



MESOPONERA Emery 

Mesoponera Emery, 1901, Annls Soc. ent. Belg. 45 : 43 [as a subgenus of Euponera]. Type- 
species: Ponera caffraria F. Smith, 1858, Cat. Hym. Brit. Mus. 6 : 91, by original designation. 
Mesoponera Emery; Wilson, 1958, Bull. Mus. comp. Zool. Harv. 119 : 349. [Raised to genus.] 

Mandibles triangular or elongate triangular, usually with more than eight teeth, without a 
basal pit on the dorsolateral surface. Palp formula 4,4. Clypeus longitudinally carinate, may 
project anteromedially as a short tooth. Promesonotal suture present, metanotal groove 
present and impressed. Propodeum compressed above, its dorsum considerably narrower than 
the pronotum. Petiole a thick scale. Middle and hind tibiae each with two spurs, one large 
and pectinate, the other small and simple. Extensor surfaces of middle tibiae without spinules; 
claws simple. Sculpture of fine dense puncturation. 

The West African species of Mesoponera nest in rotten wood on the ground and 
forage in the leaf-litter and in the soil below logs and stones. 

PALTOTHYREUS Mayr 

Paltothyreus Mayr, 1862, Verh. zool.-bot. Ges. Wien. 12 : 735. Type-species: Formica tarsata 
Fabricius, 1798, Ent. Syst. Suppl. : 280, by monotypy. 

The single species of this genus, P. tarsatus (F.), is the common 'Stink Ant'. 

Large, total length 15 mm or more. Clypeus with median portion abruptly raised and 
projecting forwards as a truncated lobe. Palp formula 4,4. Promesonotal suture distinct, 
metanotal groove indicated by a weak impression which does not break the sculpture. Petiole 
a thick scale. Middle and hind tibiae each with a large pectinate and a small simple spur, 
claws with a median tooth. Head and alitrunk striate dorsally, strongly arched on the pronotum ; 
Generally the sculpturation becomes finer from the front to the back of the dorsum of the 
alitrunk, disappearing on the gaster. 

Nests are made directly into the ground, either in the open or with the nest 
entrance beside a rock or piece of wood. The ants are general scavengers and carni- 
vores, usually foraging singly but occasionally found in small processions. 

PHRYNOPONERA Wheeler 

Phrynoponera Wheeler, 1920, Psyche, Camb. 27 : 53. Type-species: Bothroponera gabonensis 
E. Andre, 1892, Revue Ent. 11 : 50, by original designation. 

Mandibles with four to seven teeth; palp formula 4,4. Median portion of clypeus bluntly 
bidentate in the more common species, but not so in all species. Promesonotal suture present, 
metanotal absent. Propodeum with two blunt, dorsoventrally flattened spines or teeth. 
Petiole a thick scale, recurved over the anterior dorsum of the first gastral segment and terminat- 
ing dorsally in a comb of five teeth. Gaster not impressed between first and second segments. 

Nests in damp-rotten logs, usually well embedded in leaf-litter or soil, and often 
associated with termites. Uncommon ants, very rarely found outside the log 
containing the nest. The genus is very closely related to Bothroponera. 



ANT GENERA OF WEST AFRICA 339 

PLECTROCTENA F. Smith 

Plectroctena F. Smith, 1858, Cat. Hym. Brit. Mus. 6 : ior. Type-species : Plectroctena mandibu- 
laris F. Smith, 1858, loc. cit., by monotypy. 

Mandibles elongate, linear, weakly curved, with not more than two blunt teeth on the inner 
margin. Mandibular articulation associated with a marked excavation of the anterior margin 
of the head in front of the eye. Palp formula 2,4. Median portion of clypeus reduced, vertical; 
the frontal carinae almost overhanging the anterior margin of the head in dorsal view. Eyes 
small to minute, occasionally depigmented. Promesonotal suture distinct; metanotal groove 
absent or represented by a weak impression on the dorsum of the alitrunk. Petiole a distinct 
node; gaster strongly impressed between first and second segments. Middle and hind tibiae 
each with a single pectinate spur; claws simple. 

Nests are made in rotten wood, usually of fallen trees with the bark still attached, 
by the larger black species of the P. mandibular is F. Smith group of species. The 
smaller, depigmented (usually red-brown) species of the P. subterranea Arnold group 
prefer wood in a much more advanced state of decay, or nest directly into hard- 
packed earth. Sculpturation throughout the genus is mostly of puncturation, 
which tends to be more coarse in the P. subterranea group, and in which the eyes 
are much more reduced than in the larger species. 

PROMYOPIAS Santschi 

Promyopias Santschi, 1914, Boll. Lab. Zool. gen. agr. Portici 8 : 323 [as a subgenus of Myopias]. 

Type-species: Myopias {Promyopias) silvestrii Santschi, 1914, op. cit.: 324, by monotypy. 
Promyopias Santschi; Emery, 1915, Boll. Lab. Zool. gen. agr. Portici 10 : 26. [Raised to genus.] 

Mandibles elongate, weakly curved, armed apical ly with three teeth, the inner margins 
retaining traces of small teeth. Median portion of clypeus not wholly vertical, the frontal 
carinae not overhanging the anterior margin of the head. Mandibular articulations not assoc- 
iated with marked excavations of the anterior margin of the head. Extensor surfaces of middle 
tibiae with numerous spines and stiff setae. Hind tibiae with two spurs, one large and pectinate, 
the other small and simple ; claws simple. Petiole a node, gaster very weakly impressed between 
first and second segments. 

The single species P. silvestrii Santschi is rare and apparently restricted to the 
forest zone. 



PSALIDOMYRMEX E. Andre" 

Psalidomyrmex E. Andre, 1890, Revue Ent. 9 : 313. Type-species: Psalidomyrmex foveolatus 
E. Andr6, 1890, op. cit. : 314, by monotypy. 

Rare, medium to large ants restricted to the forest zones of West and Central 
Africa. 

Mandibles edentate to weakly toothed, varying in shape from subtriangular to falcate, 
always with the apex prolonged into a long, acute point, crossing over at rest. Basal mandibular 
groove distinct. Palp formula 3,4. Promesonotal suture distinct, mobile, metanotal absent. 
Petiole a node ; gaster strongly impressed between first and second segments. Middle and hind 
tibiae with a single pectinate spur, claws simple. Basic sculpture consists of large shallow 
foveolae or pits, from each of which a single seta arises. The interspaces between such pits 
usually finely striate. 



34© B. BOLTON 

The species of this uncommon genus nest in rotten wood, usually in an advanced 
state of decay, and forage in and below rotten logs and in deep leaf-litter. 

TRACHYMESOPUS Emery 

Trachymesopus Emery, 1911, Genera Insect., Ponerinae : 84 [as a subgenus of Euponerd). 

Type-species: Formica stigma Fabricius, 1804, Syst. Piez. : 400, by original designation. 
Trachymesopus Emery; Wilson, 1958, Bull. Mus. comp. Zool. Harv. 119 : 352. [Raised to 

genus.] 

Mandibles triangular, dentate; basally with a distinct pit or fovea on the dorsolateral surface. 
Palp formula 4,4. Clypeus carinate medially. Eyes small, their maximum diameter less than 
the maximum width of the scape. Promesonotal suture distinct, metanotal groove present, 
very weakly impressed. Propodeum compressed, its dorsum narrower than that of the pro- 
notum. Petiole a thick scale ; gaster distinctly impressed between the first and second segments. 
Middle and hind tibiae each with one large pectinate and one small simple spur, claws simple. 
Sculpture of fine dense puncturation. 

The single species occurring in West Africa, T. brunoi (Forel), is quite common in 
wooded areas, nesting either in rotten wood or in soil and foraging in leaf-litter, 
rotten wood, or in the earth, often being found with termites. 

Subfamily CERAPACHYINAE 
Tribe CERAPACHYINI 
CERAPACHYS F. Smith 

Cerapachys F. Smith, 1857, /. Proc. Linn. Soc. 2 : 74. Type-species: Cerapachys antennatus 
F. Smith, 1857, loc. cit., by monotypy. 

Antennae 11- or 12- segmented, the apical funicular segment greatly swollen and forming 
a one-segmented club. Mandibles edentate or denticulate; palp formula 2,2 (Gotwald, 1969.) 
Frontal carinae raised, exposing the condylar bulbs of the antennae in dorsal view. Genae 
longitudinally carinate ; eyes present. Dorsum of alitrunk devoid of sutures. Petiole a distinct 
and massive node; gastral constriction between the first and second segments often extreme, 
so that the pedicel in some species consists, to all intents and purposes of petiole and postpetiole. 
Petiole never marginate laterally. Pygidium impressed, armed laterally or posteriorly with a 
row of spines or denticulae. Middle and hind tibiae with two spurs, claws simple. 

Usually small, black or brown-black ants nesting in soil, rotten twigs or small 
pieces of wood in the leaf-litter. Specimens are most often obtained by use of a 
Berlese funnel. Nests may occasionally be found under the bark of trees, at or near 
ground level. Uncommon ants, all known species raid the nests of other ants for 
food. 

PHYRACACES Emery 

Phyracaces Emery, 1902, Re. Sess. Accad. Sci. 1st. Bologna, N.S. 6 : 23. Type-species: Cera- 
pachys mayri Forel, 1892, in Grandidier, Hist. Nat. Phys. Madagascar 20 : 244, by original 
designation. 
Closely related to, and answering to the description of Cerapachys above; differing in the 

following respects : 



ANT GENERA OF WEST AFRICA 341 

Antennae 12-segmented, the apical three funicular segments forming a club. Palp formula 
4,3 in species examined. Node of petiole distinctly marginate laterally and often armed 
posterodorsally with a pair of teeth. 

Nesting and foraging as in Cerapachys. 

SPHINCTOMYRMEX Mayr 

Sphinctomyrmex Mayr, 1866, Verh. zool.-bot. Ges. Wien 16 : 895. Type-species : Sphinctomyrmex 

stahli Mayr, 1866, loc. cit., by monotypy. 
Aethiopopone Santschi, 1930, Bull. Annls Soc. ent. Belg. 70 : 49. Type-species: Sphinctomyrmex 

rufiventris Santschi, 1915, Annls Soc. ent. Fr. 84 : 244, by original designation. 

Answering to the description of Cerapachys above but differing in the following respects: 
Antennae of twelve segments, club of funiculus not formed of apical segment only. Palp 

formula 3,3 (Gotwald, 1969). Eyes absent. Segments of gaster separated from each other 

by distinct constrictions. 

Rare, collected only from Berlese funnel extracts of forest leaf-litter. 

Tribe CYLINDROMYRMICINI 

SIMOPONE Forel 

Simopone Forel, 1891, in Grandidier, Hist. Phys. Nat. Madagascar 20, 2 : 139. Type-species: 
Simopone grandidieri Forel, 1891, op. cit.: 141, by monotypy. 

Mandibles edentate or weakly denticulate. Palp formula 6,4 or 3,2 in species examined. 
Frontal carinae reduced, but not usually vertical, the condylar bulbs of the antennae visible. 
Antennae 11-segmented, the scape short, flattened and subtriangular in shape. Eyes large, 
ocelli present. Dorsum of alitrunk devoid of sutures although the line of the promesonotal 
suture may be marked by a row of pits or short longitudinal ribs. Petiole a low node; con- 
striction between first and second gastral segments deep. Pygidium impressed, armed on each 
side by a row of stumpy spines. Middle tibiae without spurs, hind tibiae each with a large 
pectinate spur; claws toothed. 

Arboreal ants, usually black in colour, more rarely depigmented and yellowish. 
Nests are made in hollow twigs or rotten branches of trees, often a considerable dis- 
tance above ground. The species are rare and appear to be mainly nocturnal in 
behaviour. 



Subfamily MYRMICINAE 
Tribe CARDIOCONDYLINI 
CARDIOCONDYLA Emery 

Cardiocondyla Emery, 1869, Annali Accad. Aspir. Nat. Napoli (2) 2 : 20. Type-species: 

Cardiocondyla elegans Emery, 1869, op. cit. : 21, by monotypy. 
Emeryia Forel, 1890, Annls Soc. ent. Belg. 34 : no. Type-species: Emeryia wroughtoni Forel, 

1890, loc. cit., by monotypy. 

Lateral portions of clypeus projecting forwards over the basal margins of the mandibles. 
Eyes well developed, situated well in front of the midlength of the head. Antennae 12-seg- 
mented, with a three-segmented club, the scapes short, failing to reach the posterior margin of 
the head. Promesonotal suture absent, metanotal groove impressed. Propodeum armed with 



342 B. BOLTON 

a pair of teeth or spines. Petiole with a long peduncle in front; in dorsal view the postpetiole 
very broad, almost or quite twice as broad as the petiole. Middle and hind tibiae without 
spurs. Sculpture usually of fine dense puncturation on head and alitrunk, absent from the 
gaster. A sparse pubescence present but no long setae except on the anterior clypeal margin 

Small to minute ants which nest in soil, usually at the bases of trees, or in com- 
pressed leaf-litter. The workers forage in the leaf-litter or ascend trees to tend 
aphids and coccids. 

Tribe CATAULACINI 

CATAULACUS F. Smith 

Cataulacus F. Smith, 1853, Trans ent. Soc. Lond. 2 : 225. Type-species: Cataulacus taprobanae 
F. Smith, 1853, loc. cit., by designation of Bingham, 1903. 

Mandibles edentate to weakly denticulate. Palp formula 5,3. Antennae n-segmented with 
a club of three segments. Antennal scrobes present, running below the eyes which are usually 
well developed and situated behind the midlength of the head. Sides of head between eye and 
occipital corner often armed with numerous laterally projecting denticles, the occipital corner 
itself usually armed with a tooth. Dorsum of alitrunk without sutures or with the sutures 
weakly marked. Sides of pronotum strongly marginate, armed with a number of small teeth or 
an angular projection. Propodeum with a pair of spines or teeth. Entire visible dorsum of 
gaster occupied by the tergite of the first segment. Head, alitrunk and gaster considerably 
dorsoventrally compressed. 

Armoured, usually black, arboreal ants nesting in hollow or rotten twigs and 
branches which are still attached to the tree. 



Tribe CREMATOGASTRINI 
CREMATOGASTER Lund 

Crematogaster Lund, 1831, Annls Sci. nat. 23 : 132. Type-species: Formica scutellaris Olivier, 
1791, Encyl. Mithod. Histoire Naturelle. Insectes 6 : 497, by designation of Bingham, 1903. 

Acrocoelia Mayr, 1852, Verh. zool.-bot. Ges. Wien. 2 : 146. Type-species: Acrocoelia ruficeps 
Mayr, 1852, op. cit. : 147 [= Formica scutellaris Olivier, 1791]. 

Tranopeltoides Wheeler, 1922, Am. Mus. Novit. no. 48 : 10. Type-species: Tranopella huberi 
Forel, 1907, Hamburg. Jb. wiss. Anst. 24 (2) : 5, by original designation. 

Mandibles with four or five teeth; palp formula usually 5,3, rarely 4,3. Antennae 10- or 11- 
segmented, the club undifferentiated or of two, three or four segments. Eyes present, usually 
well developed, situated at or just behind the midlength of the side of the head. Promesonotal 
suture usually represented by a weak impression which fails to break the sculpture, more 
rarely it is well developed or absent. Metanotal groove impressed, often deeply so. Propodeum 
armed with a pair of spines or teeth, rarely reduced to tubercles or absent. Petiole dorso- 
ventrally flattened, without a node, the dorsal surface weakly convex to weakly concave. 
Postpetiole a node, often with a median longitudinal groove, attached to the dorsum of the first 
gastral segment. Gaster heart-shaped or triangular in dorsal view. Sting spatulate. Tarsal 
claws simple, often large. 

A large genus of mostly arboreal ants. In the majority of cases nests are made in 
hollow or rotten twigs and branches, or directly into the trunk of the tree, but in 
some cases a large carton-nest of chewed wood is made and attached to the trunk, 
often high up the tree. 



ANT GENERA OF WEST AFRICA 343 

The unusual construction of the pedicel allows the gaster to be reflexed over the 
alitrunk, and when agitated the ants run about with the gaster held in this position. 
Most species are avid tenders of coccids and some build protective tents of vegetable 
matter over aggregations of coccids. 

Tribe DACETINI 
CODIOMYRMEX Wheeler 

Codiomyrmex Wheeler, 1916, Bull. Mus. comp. Zool. Harv. 60 : 326. Type-species: Codio- 
myrmex thaxteri Wheeler, 1916, op. cit., : 327, by original designation. 

Mandibles short, subtriangular, with less than ten teeth arranged in such a way that in dorsal 
view only three or four are visible, the rest arranged in a more or less vertical series on the 
down-curved apical portion of the mandible. Antennae six-segmented, the second and third 
funicular segments reduced. Antennal scrobes present above the eyes. Dorsum of alitrunk 
without sutures; propodeum armed with a pair of small teeth subtended by a lamella running 
the height of the declivity. Petiole and postpetiole with well developed spongiform appendages. 

Clypeus smooth and shining with abundant simple setae, rest of head reticulate-punctate. 
Dorsum of alitrunk, postpetiole and first gastral segment mostly smooth and shiny, the last with 
distinct longitudinal striations. 

Minute (2 mm or less) brown-black ants found as yet only in Berlese funnel samples 
of forest leaf-litter in West Africa. 

EPITRITUS Emery 

Epitritus Emery, 1869, Boll. Soc. ent. Ital. 1 : 136. Type-species: Epitritus argiolus Emery, 
1869, loc. cit., by monotypy. 

Mandibles elongate with 0-1 preapical teeth. Apical armament consisting of a long dorsal 
spiniform tooth subtended by a vertical series of denticulae or by the denticulae alone. Labrum 
projecting anteriorly between the mandibles as a biconical structure. Antennae 4- or 6- 
segmented, the scape lobiform basally. Antennal scrobes present above the reduced eyes. 
Promesonotal suture represented by a weak impression or absent, similarly with the metanotal 
groove, which is however usually more distinct than the promesonotal suture. Spongiform 
appendages of pedicel greatly reduced or absent. Head with numerous large orbicular hairs. 
Sculpture finely reticulate-punctate. 

Minute ants, 1-2-2-2 mm total length. 

Three West African species are known, one from the savannah region of Northern 
Nigeria, the other two from Eastern Ghana, in the forest zone. The savannah 
species nests in rotten wood which is probably true of the forest forms, known at 
present only from Berlese funnel samples of leaf-litter. 

MICCOSTRUMA Brown 

Miccostruma Brown, 1948, Trans Am. ent. Soc. 74 : 123. Type-species: Epitritus mandibulars 
Szabo, 1909, Archvm zool., Bpcst 1 (7) : 1-2, by original designation. 

Mandibles subtriangular, serially dentate, concealed for most of their length by the very 
well developed, anteriorly projecting clypeus, which is fringed anteriorly and laterally by 
flattened hairs. Antennae 4-segmented; antennal scrobes present. Sutures absent from 
dorsal alitrunk. Propodeum bidentate, with ventral laminae running down the declivity. 



344 B. BOLTON 

Spongiform appendages of pedicel well developed. Dorsum of head with fine, scattered punc- 
tures, hairs absent except on clypeus. Dorsum of alitrunk, pedicel and gaster mostly shiny 
with small, widely spaced punctures. Base of first gastric tergite with longitudinal striae. 

Minute ants, about 2 mm total length. Foraging workers occur in leaf-litter. 



MICRODACETON Santschi 

Micvodaceton Santschi, 1913, Bull. Soc. ent. Fr. : 478. Type-species: Microdaceton exornatum 
Santschi, 1913, loc. cit., by monotypy. 

Mandibles elongate, linear, without preapical armament; apical armament of three spini- 
form teeth. Antennae 6-segmented; antennal scrobes absent. Gena in front of eye with a 
laterally projecting tooth; occipital lobes of head armed with teeth. Promesonotum poster- 
iorly, propodeum and petiole bispinose or bidentate; postpetiole with lateral alar appendages. 

Uncommon. Recovered from leaf-litter by Berlese funnel extraction and also 
from soil-pockets on trees. 



QUADRISTR UMA Brown 

Quadristruma Brown, 1949, Trans. Am. ent. Soc. 75 : 47. Type-species: Epitritus emmae 
Emery, 1890, Boll. Soc. ent. Ital. 22 : 70, by original designation. 

Mandibles elongate, linear, strongly curved; preapical armament a single long, spiniform 
tooth, apical armament a fork of two teeth. Labrum not projecting between mandibles as a 
biconical structure. Antennae 4-segmented; antennal scrobes present. Head with orbicular 
hairs present. Sculpture a fine puncto-reticulation, first gastral segment shiny except for 
basigastric costulae. 

Q. emmae (Emery), a pantropical tramp species, is here recorded for the first time 
from West Africa. A single alate female was taken on a sticky trap hung from a 
cocoa tree (Theobroma cacao L.) at Bunso, Eastern Region of Ghana, in early October, 
1969. Previously Brown (1954) had indicated that Quadristruma was most likely 
of African origin and its presence on the continent is now confirmed. 



SERRASTRUMA Brown 

Serrastruma Brown, 1948, Trans. Am. ent. Soc. 74 : 107 [as a subgenus of Smithistruma). Type- 
species: Strumigenys simoni Emery, 1895, Annls Soc. ent. Fr. 63 : 42, by original designation. 
Serrastruma Brown; Brown, 1949, Mushi 20 : 6. [Raised to genus.] 

Mandibles elongate triangular, serially denticulate, with more than twenty denticulae, the 
basal 4-8 of which may be much larger than those preceding. Antennae 6-segmented, the 
second and third segments of the funiculus reduced; antennal scrobes present. Promesonotal 
suture represented by a weak impression or absent. Sometimes pronotum separated from 
mesonotum by a change in sculpture. Metanotal groove impressed; propodeum armed with 
a pair of teeth, may be reduced. Spongiform appendages of pedicel usually reduced. 

Small yellow-brown ants nesting in rotten wood and foraging there and in leaf- 
litter. One species, S. maynei (Forel), is subarboreal, nesting in small rot-holes in 
the trunks and branches of low trees. This genus and Strumigenys constitute 
the most common members of the tribe Dacetini in West Africa. 



ANT GENERA OF WEST AFRICA 345 

SMITHISTRUMA Brown 

Smithistrnma Brown, 1948, Trans Am. ent. Soc. 74 : 104. Type-species: Strumigenys pulchella 

Emery, 1895, Zool. Jb. Syst. 8 : 327, by original designation. 
Cephaloxys F. Smith, 1864, /. Proc. Linn. Soc. 8 : 76, nom. preocc. 

Mandibles triangular, armed with 10 or less small teeth or denticulae. Antennae 6-segmented, 
the second and third funciular segments reduced, may be very small; antennal scrobes present. 
Head often very much elongated. Promesonotal suture absent or marked by a weak impres- 
sion; metanotal groove weakly or not impressed. Spongiform appendages of pedicel usually 
well developed. 

Uncommon, nesting in rotten twigs or in compressed leaf-litter, occasionally 
directly into soil at the bases of trees. Most workers are retrieved from Berlese 
funnel samples of leaf-litter. 

STRUMIGENYS F. Smith 

Strumigenys F. Smith, i860, /. ent. London 1 : 72. Type-species: Strumigenys mandibularis 

F. Smith, i860, loc. cit., by monotypy. 
Labidogenys Roger, 1862, Berl. ent. Z. 6 : 249. Type-species: Labidogenys lyroessa Roger, 

1862, op. cit. : 251, by monotypy. 
Pyramica Roger, 1862, Berl. ent. Z. 6 : 251. Type-species: Pyramica gundlachi Roger, 1862, 

op. cit. : 253, by monotypy. 
Proscopomyrmex Patrizi, 1946, Boll. 1st. ent. Univ. Bologna 15 : 294. Type-species: Proscopo- 

myrmex londianensis Patrizi, 1946, op. cit., : 295, by original designation. 
Eneria Donisthorpe, 1948, Ann. Mag. nat. Hist. (11) 14 : 598. Type-species: Eneria excisa 

Donisthorpe, 1948, loc. cit., by original designation. 

Mandibles elongate, linear; preapical armament usually of two teeth on the distal portion of 
the blade, occasionally with one tooth missing from one of the blades. Apical armament of 
a fork of two spiniform teeth, with or without intercalary denticles. Antennae 6-segmented, 
the second and third funicular segments reduced, sometimes so reduced that the antennae 
appears four-segmented. Antennal scrobes present. Often the ventrolateral margin of the 
head is excised in front of the eye forming the characteristic preocular notch of S. rogeri Emery 
and allies. 

Nests in rotten wood, leaf-litter, or directly into hard-packed earth. Two species 
are arboreal and nest in rot-holes in the trunks and branches of low trees. The 
smaller species often nest in small twigs in the leaf-litter or in compressed leaf- 
mould and are quite common in Berlese funnel samples from the forest zone. 

TRICHOSCAPA Emery 

Trichoscapa Emery, 1869, Annali Accad. Aspir. Nat. Napoli (2) 2 : 24 [as a subgenus of Strumi- 
genys]. Type-species: Strumigenys (Trichoscapa) membranifera Emery, 1869, loc. cit., by 
monotypy. 

Trichoscapa Emery; Brown, 1948, Trans Am. ent. Soc. 74 : 11 2-1 14. [Raised to genus.] 

Mandibles triangular with strong horizontal basal borders which are not covered by the 
clypeus at full closure. Antennae 6-segmented; antennal scrobes present. Head devoid of 
hairs except for two on the posterodorsal surface and five or six on each antennal scape. Sides 
of pronotum strongly marginate. 

A tramp ant spread by human commerce, T. membranifera Emery occurs in many 
parts of the tropics and warm temperate zones. Brown (1949a) states that the 



346 B. BOLTON 

species is probably of African origin, but the species has not yet been recorded 
from Africa. Wilson & Taylor (1967) report that 'The species has an ecological 
amplitude unusual for a dacetine, nesting in major habitats from dense woodland 
to dry, open cultivated fields.' In view of the above, Trichoscapa is included in the 
present work. 



Tribe LEPTOTHORACINI 
ADELOMYRMEX Emery 

Adelomyrmex Emery, 1897, Termeszetr. Fuz. 20 : 590. Type-species: Adelomyrmex biroi 
Emery, 1897, loc. cit., by monotypy. 

Mandibles subtriangular, dentate and with a single tooth on the basal internal margin. 
Maxillary palp with a single segment (Gotwald, 1969). Median portion of clypeus projecting 
over the basal borders or the mandibles when fully closed. Antennae 12-segmented with a 
funicular club of two segments, of which the apical segment is much the larger. Eyes small, 
situated just anterior to the midline of the side of the head. Promesonotal suture absent; 
metanotal groove impressed. Propodeum armed with a pair of spines. 

Recovered from Berlese funnel samples of leaf-litter from the forest zone, it is 
probable that this genus has been spread by human commerce from the Papuan 
region to West Africa. 



LEPTOTHORAX Mayr 

Leptothorax Mayr, 1855, Verh. zool.-bot. Ges. Wien 5 : 431. Type-species: Formica acervoritm 

Fabricius, 1804, Syst. Piez. : 407, by designation of Wheeler, 191 1. 
Goniothorax Emery, 1896, Boll. Soc. ent. Ital. 28 : 26 [as a subgenus of Leptothorax], nom. 

preocc. 
Caulomyrma Forel, 1914, Bull. Soc. vaud. Sci. nat. 50 : 233 [as a subgenus of Leptothorax}. 

Type-species: Leptothorax echinatinodis Forel, 1886, Annls. Soc. ent. Belg. 30 : 48, by original 

designation. 
Limnomyrmex Arnold, 1948, Occ. Pap. natn. Mus. Sth. Rhodesia 2 (14) : 222. Type-species: 

Limnomyrmex stramineus Arnold, 1948, op. cit. : 223, by original designation. 

Median portion of clypeus projecting over basal borders of mandibles as a broad, arcuate 
lobe; the posterior margins of the lateral portions of the clypeus not raised into a ridge in 
front of the antennal insertions. Antennae 12-segmented with a 3-segmented club; antennal 
scrobes absent. Eyes well developed, situated at about the midlength of the side of the head. 
Anterodorsal pronotal angles acute, giving a square shouldered appearance in dorsal view. 
Promesonotal suture absent, metanotal groove absent to weakly impressed. Propodeum 
bidentate or bispinose. Petiole with a very short, thick anterior peduncle. Erect setae on all 
dorsal surfaces of body short, thick and blunt. 

Nests in rotten wood, usually embedded in leaf-litter. Foragers are found in 
leaf-litter, log-mould, or more rarely running about on open ground. 

Superficially resembling some species of Tetramorium, Leptothorax may be dis- 
tinguished by a combination of the characters noted above and by the fact that in 
Tetramorium the sting is equipped apicodorsally with a translucent, triangular 
lamelliform appendage, absent in Leptothorax. 



ANT GENERA OF WEST AFRICA 347 

MACROMISCHOIDES Wheeler 

Macromischoid.es Wheeler, 1920, Psyche, Camb. 27 : 53. Type-species: Macromischa aculeata 
Mayr, 1866, Sber. Akad. Wiss. Wien 53(i) : 507, by original designation. 

Mandibles subtriangular with 9 to 1 1 teeth of varying size. Palp formula 3,2. Head narrowed 
in front, broadening behind to the large, protuberant eyes and narrowing posterior to the eyes. 
Antennae 12-segmented with a club of three segments; the scape very long, easily surpassing 
the occipital margin. Pronotum with a small, obtuse tubercle at each side. Promesonotal 
suture absent; metanotal groove impressed. Propodeum with a pair of long acute spines. 
Petiole with a long narrow anterior peduncle. Body with numerous long, fine pointed setae. 

Arboreal, constructing nests of vegetable fragments attached to the ventral 
surfaces of leaves or in the axils of larger leaves. Predacious, principally nocturnal 
ants. 



Tribe MELISSOTARSINI 
MELISSOTARSUS Emery 

Melissotarsus Emery, 1877, Annali Mus. civ. Stor. nat. Giacomo Doria. 9 : 378. Type-species: 
Melissotarsus beccarii Emery, 1877, op. cit. : 379, by monotypy. 

Antennae 6-segmented with a two-segmented club, scapes very short, extending about half 
the distance from their insertions to the occipital margin. Antennal scrobes absent. Eyes well 
developed, situated anterolaterally and noticeably longer than broad. Dorsum of alitrunk 
devoid of sutures, smoothly convex in profile; propodeum unarmed. Postpetiole very broadly 
attached to gaster posteriorly. Coxae of middle and hind legs very large, much larger than 
fore coxae. Basal (first) segment of tarsi enlarged, as broad as or broader than the tibiae; 
remaining tarsal segments small. 

These uncommon ants nest under the bark of trees, usually some distance above 
the ground. They are sluggish and often feign death when disturbed, and are only 
rarely seen outside the nest during the day. 

Tribe MERANOPLINI 
CALYPTOMYRMEX Emery 

Calyptomyrmex Emery, 1887, Annali Mus. civ. Stor. nat. Giacomo Doria. 25 : 471. Type- 
species: Calyptomyrmex beccarii Emery, 1887, op. cit. : 472, by monotypy. 

Weberidris Donisthorpe, 1948, Entomologist's mon. Mag. 84 : 281. Type-species: Weberidris 
rufobrunnea Donisthorpe, 1948, loc. cit., by original designation. 

Median portion of clypeus vertical, projecting anterodorsally as a bilobed structure, over- 
hanging the mandibles. Antennae 11- or 12-segmented, with a three-segmented club. An- 
tennal scrobes present above the eyes, very deep and capable of containing the whole antenna. 
Dorsum of alitrunk devoid of sutures, promesonotum convex in profile, the propodeum sloping 
(sometimes steeply) and armed with a pair of spines or teeth. Dorsal surfaces of body usually 
with spatulate or otherwise bizarre setae. 

Cryptic species, usually nesting in very rotten wood deeply embedded in the 
ground, or in compressed leaf-litter. Foragers are found in log- and leaf-mould and 
occasionally in the soil beneath rotten twigs and stones. 



348 B. BOLTON 

MERANOPLUS F. Smith 

Meranoplus F. Smith, 1854, Trans ent. Soc. Lond. (2) 2 : 224. Type-species: Cryptocerus bicolor 
Guerin, 1845, Iconog. Regn. Anim. 7 : 425, by designation of Bingham, 1903. 

Cryptocephalus Lowne, 1865, Entomologist 2 : 336. Type-species: Cryptocerus pubescens 
F. Smith, 1853, Trans ent. Soc. Lond. (2) 2 : 223, by monotypy. 

Antennae 9-segmented with a 3-segmented club. Antennal scrobes present, deep; the eyes 
well developed and situated towards the posterior end of the scrobe on its ventral border. 
Promesonotum fused, strongly margined all round and projecting posteriorly as a shelf, over- 
hanging the propodeum which is vertical or nearly so. Posterior margin of the promesonotal 
shelf armed with a number of teeth or spines. Propodeum unarmed or bidentate or bispinose. 
Petiole and postpetiole squamiform. Head, alitrunk and gaster clothed with abundant long, 
soft setae, giving the ant a furry appearance to the naked eye. 

The forest species usually nest directly into the ground or amongst the roots of 
low plants, and forage in the leaf-litter and below rotten logs. Savannah species 
adopt similar nesting sites but are much more active in the open and are often to 
be seen running on termite mounds. 

Tribe MYRMECININI 

ATOPOMYRMEX E. Andre 

Atopomyrmex E. Andre, 1889, Revue Ent. 8 : 226. Type-species: Atopomyrmex mocquerysi 
E. Andre, 1889, op. cit. : 227, by monotypy. 

Polymorphic. Palp formula 4,3. Antennae 12-segmented with a club of three segments. 
Antennal scrobes absent. Eyes well developed, flat, situated on the posterior half of the 
sides of the head. Pronotum marginate laterally, the promesonotal suture absent. Mesonotum 
bituberculate behind. Metanotal groove impressed; propodeum armed with a pair of long 
spines. Petiole armed with a pair of spines above. 

Arboreal ants, nesting in the wood of large trees, usually a considerable distance 
above the ground. 

PRISTOMYRMEX Mayr 

Pristomyrmex Mayr, 1866, Verh. zool.-bot. Ges. Wien 16 : 903. Type-species: Pristomyrmex 
pungens Mayr, 1866, op. cit. : 904, by monotypy. 

Odontomyrmex E. Andre, 1905, Revue Ent. 24 : 207 [as a subgenus of Pristomyrmex]. Type- 
species: Pristomyrmex (Odontomyrmex) quadridentatus E. Andre, 1905, loc. cit., by monotypy. 

Hylidris Weber, 1941, Ann. ent. Soc. Am. 34 : 190. Type-species: Hylidris myersi Weber 
1941, loc. cit., by original designation. 

Dodous Donisthorpe, 1946, Proc. R. ent. Soc. Lond. (B)15 : 145. Type-species: Dodous tri- 
spinosus Donisthorpe, 1946, loc. cit., by original designation. 

Palp formula 1,3 or 2,3. Median portion of clypeus with a number of blunt teeth on the 
anterior margin; lateral portions reduced to a thin raised strip in front of the antennal insertions. 
Antennae 1 i-segmented with a 3-segmented club. Eyes medium or small, situated at about the 
midlength of the head ; the head itself broader in front than behind, the sides convex. Dorsum 
of alitrunk devoid of sutures. Pronotum usually armed with a pair of spines of blunt tubercles, 
very rarely unarmed. Propodeum bispinose or bidentate. Sting simple, without a spatulate 
appendage. 

Nests in rotten wood or fallen twigs in the leaf -litter; foraging is carried out in 
the leaf-litter or beneath the bark of fallen trees. This genus may be confused 



ANT GENERA OF WEST AFRICA 349 

with some species of the genus Xiphomyrmex which are superficially similar but in 
Xiphomyrmcx the clypeal margin and pronotum are unarmed, and the sting has a 
spatulate apical portion. 

TERATANER Emery 

Terataner Emery, 1912, A mils Soc. ent. lielg. 61 : 103. Type-species: Atopomyrmex forcli 

Emery, 1899, Boll. Soc. ent. Ital. 31 : 274, by original designation. 

Antennae 1 '-segmented with a three-segmented club Antennal scrobes absent. Palp 
formula 4,3. Pronotum anteriorly, and promesonotum laterally marginate. Promesonotal 
suture absent but lateral marginatum broken at junction of pro- and mesonotum. Metanotal 
groove weakly impressed; propodeum unarmed, metapleural lobes present. Petiole with a 
pair ol teeth above. 

Arboreal, nesting in the trunk or branches of living trees ; foraging wholly arboreal, 
the ants of this genus arc never found on the ground under normal circumstances. 

Tribe MYRMICARIINI 

MYRMICARIA Saunders 

Myrmicaria Saunders, 1841, Trans cut. Soc. Loud. 3 : 57. Type-species: Myrmicaria brunnea 

Saunders, 1841, loc. cit., by monotypy. 
Heptacondylus P. Smith, 1857, J. Pvoc. Linn. Soc. 2 : 71. Type-species: Heptacondylus ;ub 

carinatus F. Smith, 1857, op. cit.: 73, by designation of Wheeler, 101 [. 
Physatta F. Smith, 1857, /. Proc. Linn. Soc. 2 : 77. Type-species: Physatta dromedarius F. 

Smith, 1.^57, op. cit.: 78, by monotypy. 

Palp formula >,,x Antennae 7-segmented, indistinctly clubbed. Frontal carinae widely 
separated; antennal scrobes absent. Eyes placed behind midlength of head. Anterolateral 
angles of pronotum drawn out into ,1 tooth on each side. Promesonotal suture represented by 
a weak impression which does not break the sculpture. Mesonotum bluntly bituberculate 
behind, sharply angled, the posterior portion more or less vertical. .Metanotal groove deeply 
impressed; propodeum bispinose. Petiole with a long anterior peduncle. Sting coarse, 
somewhat flattened from side to side. Head, body and appendages with numerous long, 
rsc, darkly coloured setae. 

Mostly savannah species. Nests are made directly into the earth, often with 
long sunken runways visible on the surface of the ground. The ants are active 
in brightest sunlight and aie general predators and scavengers. In life the peduncle 
of the petiole is held almost vertically so that the gaster points almost straight down 
from base to apex, giving the ant a peculiarly foreshortened appearance. 

Tribe MYRMICINI 
CRATOMYRMEX Emery 

Cratomyrmex Emery, 1891, Annls Soc. ent. Fr. 60 : 572. Type-species: Cratomyrmex regalis 

Emery, 1819, loc. cit., by monotypy. 

Answering to the description of Messor below but with the tibial spurs of the middle and hind 
legs pectinate. 

The genus is confined to savannah regions, having the same nesting sites and 
habits as Messor. 



35© B. BOLTON 

Tribe OCHETOMYRMICINI 

WASMANNIA Forel 

Wasmannia Forel, 1893, Trans ent. Soc. Lond. : 383. Type-species: Tetramoriwn auropitnct- 
atum Roger, 1863, Berl. ent. Z. 7 : 182, by designation of Wheeler, 191 1. 

Hercynia Enzmann, 1947, Jl N.Y. ent. Soc. 55 : 43. Type-species: Hercynia panamana Enz- 
mann, 1947, op. cit. : 44, by original designation. 

Minute ants, 2 mm or less in total length. Palp formula 3,2 (Gotwald, 1969). Antennae 
11-segmented with a three-segmented club. Antennal scrobes present, bounded below by a 
weak longitudinal carina running above the eye. Anterodorsal angles of the pronotum acute; 
pronotum strongly marginate anteriorly. Promesonotal suture absent; metanotal groove 
weakly impressed. Propodeum bispinose; metapleural lobes present. 

A single species, W. auropunctata (Roger,) of this neotropical genus has been intro- 
duced into Cameroun, where in places it is quite successful. It has not yet been 
reported as being successfully introduced into West Africa. The minute workers 
forage arboreally and tend aphids and coccids as well as being active predators. 

Tribe PHEIDOLINI 

MESSOR Forel 

Messov Forel, 1890, Annls Soc. ent. Belg. 34 C.R. : 68 [as a subgenus of Aphaenogaster]. Type- 
species: Formica barbara Linnaeus, 1767, Syst. Nat. ed. 12,2 : 962, by designation of Bingham 
1903. 

Messor Forel; Emery, 1908, Dt. ent. Z. : 437. [Raised to genus.] 

Polymorphic. Mandibles strongly curved, usually dentate in smaller workers, more or less 
edentate in larger. Head as broad as or broader then long; eyes situated in the middle of the 
sides of the head. Antennae 12-segmented, without a differentiated club. Gular surface of 
head with a psammophore. Promesonotal suture present; metanotal groove impressed. Pro- 
notum and mesonotum together dome-shaped in profile, the propodeum more or less flat and 
considerably depressed, on a much lower level than the apex of the promesonotal 'dome,. 
Propodeum with a pair of blunt teeth. Node of petiole emarginate above in large workers, 
this character may be reduced or absent in small workers. Spurs of middle and hind tibiae 
simple. 

Restricted to the savannah regions of West Africa and also occurring on the 
coastal plains. Nests are made directly into the earth in open ground, and have a 
crater-like entrance. 



PHEW OLE Westwood 

Pheidole Westwood, 1840, Ann. Mag. nat. Hist. 6 : 87. Type-species: Atta providens Sykes, 
1835, Trans, ent. Soc. Lond. 1 : 103, by designation of Bingham, 1903. 

Leptomyrma Motschulsky, 1863, Bull. Soc. Nat. Moscou 36 : 17. Type-species: Leptomyrma 
gracilipes Motschulsky, 1863, loc. cit., by monotypy. 

Epipheidole Wheeler 1903, Bull. Am. Mus. nat. Hist. 19 : 664. Type-species: Epipheidole 
inquilina Wheeler 1903, loc. cit., by monotypy. 

Dimorphic, rarely with intermediates. Major worker (soldier) with head massive, the occipital 
margin deeply impressed in the middle. Mandibles large, heavy, strongly curved, each usually 
armed with three teeth, of which two are situated apically and one basally on the masticatory 



ANT GENERA OF WEST AFRICA 351 

margin and separated by a diastema. Palp formula 2,2 in species examined. Antennae 12- 
segmented with a 3-segmented club. Eyes situated in front of the midlength of the side of 
the head. Promesonotal suture absent or represented by an impressed line, rarely present. 
Metanotal groove deeply impressed. Propodeum armed with a pair of spines or teeth; petiole 
usually emarginate above. 

Minor worker with occipital margin of head shallowly emarginate or more usually with the 
sides of the head converging behind the eyes so that the occipital margin is very short. Man- 
dibles usually with two or three large teeth apically followed by a row of denticulae of uneven 
sizes. Palp formula and antennae as in major worker but eyes usually set just in front of 
midlength of the head. Alitrunk and pedicel as above. 

A very common genus in West Africa, present throughout the region and nesting 
in rotten wood, directly into open ground, under stones and logs, amongst the roots 
of plants and occasionally beneath the bark of standing trees or in rot-holes in tree 
trunks. 

Tribe SOLENOPSIDINI 

CAREBARA Westwood 

Carebara Westwood, 1841, Ann. Mag. nat. Hist. 6 : 86. Type-species: Carebara lignata West- 
wood, 1841, loc. cit., by monotypy. 

Mandibles with five or six teeth. Palp formula 2,2 (Ettershank, 1966). Antennae Q-seg- 
mented with a club of two segments. Clypeus bicarinate; eyes completely absent. Promeso- 
notal suture absent; metanotal groove impressed. Propodeum unarmed. 

Workers minute, usually less than 2 mm total length; depigmented, usually yellow in colour. 

The ants of this genus are wholly hypogaeic and are nearly always found with 
termites. Their nests are usually built in the walls of termitaria of species which 
do not show a mound above ground-level. Occasional workers are found in Berlese 
funnel samples. At the times when alate females and males are produced, the 
workers emerge in hundreds onto the surface of the ground. 

DIPLOMORIUM Mayr 

Diplomorium Mayr. 1901 Annln naturh. Mus. Wien 16 : 16. Type-species: Diplomorium 
longipenne Mayr, 1901, op. cit. : 18, by monotypy. 

Bondroitia Forel, 191 1, Bull. Soc. vaud. Sci. nat. 47 : 398 [as a subgenus of Diplomorium]. Type- 
species: Diplomorium lujae Forel, 1909, Annls Soc. ent. Belg. 53 : 72, by monotypy. 

As Monomorium below but always with the antennae 11 -segmented and the median portion 
of the clypeus swollen but not bicarinate. Palp formula 2,2 (Ettershank, 1966). 

The species nest in rather dry rotten wood. 

MONOMORIUM Mayr 

Monomorium Mayr, 1855, Verh. zool.-bot. Ges. Wien 5 : 452. Type-species: Monomorium 

minutum Mayr, 1855, op. cit. : 453, by monotypy. 
Phacota Roger, 1862, Berl. ent. Z. 6 : 260. Type-species: Phacota sicheli Roger, 1862, op. cit.: 

262, by monotypy. 
Trichomyrmex Mayr, 1865, Reise der . . . Fregatte Novara, Zool. 2 (1) : 19. Wien. Type-species: 

Trichomyrmex rogeri Mayr, 1865, loc. cit., by monotypy. 

C* 



352 B. BOLTON 

Lampromyrmex Mayr, 1868, Beitr. Naturk. Preuss. 1 : 92. Type-species: Lampromyrmex 

gracillimus Mayr, 1868, nom. preocc. [= Monomorium mayrianum Wheeler, 1914, Schr. 

phys.-okon. Ges. Konigsb. 55 : 45. (fossil), nom. substit.], by monotypy. 
Holcomyrmex Mayr, 1878, Verh. zool.-bot. Ges. Wien 28 : 671. Type-species: Holcomyrmex 

scabriceps Mayr, 1878, op. cit. : 672, by designation of Bingham, 1903. 
Epoceus Emery, 1892, Annls Soc. ent. Fr. 61 : 272. Type-species: Epoecus pergandei Emery, 

1892, op. cit. : 273, by monotypy. 
Wheelevia Forel, 1905, Annls Soc. ent. Belg. 49 : 171, nom. preocc. Type-species: Wheeleria 

santschii Forel, 1905, loc. cit., by monotypy. 
Wheeleriella Forel, 1907, Int. Sci. Revue 4 : 145, nom. substit. pro. Wheeleria Forel. 
Epixenus Emery, 1908, Dt. ent. Z. : 556. Type-species: Monomorium advena Brown and 

Wilson, 1957, Ent. News 68 : 244, nom. substit. pro Epixenus andrei Emery, 1908 (nee 

Saunders, 1890). 
Xeromyrmex Emery, 1915, Bull. Soc. ent. Fr. : 190 [as a subgenus of Monomorium]. Type- 
species: Formica salomonis Linnaeus, 1758, Syst. Nat. ed. 10 : 580, by original designation. 
Parholcomyrmex Emery, 1915, Bull. Soc ent Fr. : 190 [as a subgenus of Monomorium] [ = 

Paraholcomyrmex, variant spelling]. Type-species: Myrmica gracillima F. Smith, 1S61, 

/. Proc. Linn. Soc. 5 : 34, by original designation. 
Corynomyrmex Viehmeyer, 1916, Arch. Naturgesch. 81 : 134 [as a subgenus of Monomorium]. 

Type-species: Monomorium (Corynomyrmex) hospitum Viehmeyer, 1916, loc. cit., by mono- 
typy. Provisional synonymy of Ettershank, 1966, Aust. J. Zool. 14 : 82. 
Isolcomyrmex Santschi, 1917, An. Soc. cient. argent. 84 : 296 [as a subgenus of Monomorium]. 

Type-species: Monomorium sanlschianum Ettershank, 1966, nom. substit. pro Holcomyrmex 

santschii Forel, 1907, nom. preocc. 
Paraphacota Santschi, 1919, Bull. Soc. ent. Fr. : 91. Type-species: Phacota noualhieri Emery, 

1895, Mem. R. Accad. Sci. 1st. Bologna 5 : 299, by original designation. 
Equestrimessor Santschi, 1919, Bull. Soc. ent. Fr. : 92. [as a subgenus of Monomorium] [= 

Equesimessor, variant spelling]. Type-species; Holcomyrmex chobauti Emery, 1897, Bull. 

Soc. ent. Fr. : 418, by designation of Donisthorpe, 1943, Ann. Mag. nat. Hist. (11) 10 : 644. 
Xenhyboma Santschi, 1919, Boln Soc. esp. Hist. nat. 19 : 405. Type-species: Xenhyboma 

mystes Santschi, 1919, loc. cit., by monotypy. Provisional synonymy of Ettershank, 1966, 

Aust. J. Zool. 14 : 82. 
Ireneidris Donisthorpe, 1943, Entomologist's mon. Mag. 79 : 81. Type-species: Ireneidris 

myops Donisthorpe, 1943, loc. cit. [= Monomorium talpa Emery, 1911, Nova Guinea 9 : 252], 

by original designation. 

Median portion of clypeus with two distinct longitudinal carinae, the clypeus sometimes 
concave between the carinae on the anterior margin and projecting as a pair of blunt teeth. 
Palp formula 1,2 or 2,2 (Ettershank, 1966). Eyes present. Antennae 11- or 12-segmented with 
a 3-segmented club. Promesonotal suture absent on dorsum of alitrunk; metanotal groove 
impressed. Propodeum unarmed; petiole pedunculate, the node high and rounded. 

The majority of West African species of this very common genus are unsculptured, 
smooth and shiny ants, but some have a universal fine dense puncturation or reticulo- 
puncturation. Nests are made in rotten wood, under stones or directly into the 
earth. M. pharaonis (L.) is a common house-inhabiting species. 



OLIGOMYRMEX Mayr 

Oligomyrmex Mayr, 1867, Tijdschr. Ent. 10 : no. Type-species: Oligomyrmex concinnus 

Mayr, 1867, op. cit.: in, by monotypy. 
Aeromyrma Forel, 1891, Annls Soc. ent. Belg. 35: 307. Type-species: Aeromyrma nosindambo 

Forel, 1891, op. cit. : 199, by monotypy. 



ANT GENERA OF WEST AFRICA 353 

Aneleus Emery, 1900, Termeszetr. Fiiz. 23 : 327 [as a subgenus of Pheidologeton.] Type-species: 
Solenopsis similis Mayr, 1862, Verh. zool.-bot. Ges. Wien 12 : 751, by designation of Wheeler, 
1911. 

Erebomyrma Wheeler, 1903, Biol. Bull. mar. biol. Lab. Woods Hole 4 : 138. Type-species: 
Erebomyrma longi Wheeler, 1903, op. cit. : 140, by monotypy. 

Lecanomyrma Forel, 191 3, Zool. Jb. Syst. 36 : 56 [as a subgenus of Pheidologeton]. Type- 
species: Pheidologeton (Lecanomyrma) butteli Forel, 1913, loc. cit., by designation of Emery, 
1922. 

Octella Forel, 1915, Ark. Zool. 9 : 69 [as a subgenus of Oligomyrmex]. Type-species : Oligomyrmex 
(Octella) pachycerus Forel, 1915, loc. cit., by original designation. 

Spelaeomyrmex Wheeler, 1922, Am. Mus. Novit. no. 45 : 9. Type-species: Spelaeomyrmex 
urichi Wheeler, 1922, loc. cit., by original designation. 

Hendecatella Wheeler, 1927, Boll. Lab. Zool. gen. agr. Portici 20 : 93 [as a subgenus of Oligo- 
myrmex] Type-species: Oligomyrmex (Hendecatella) capreolns Wheeler, 1927, loc. cit., 
by monotypy. 

Solenops Karawajew, 1930, Zool. Anz. 92 : 207 [as a subgenus of Solenopsis]. Type-species: 
Solenopsis (Solenops) weyeri Karawajew, 1930, loc. cit., by monotypy (nee Solenops Dufour, 
1820, Arachnida). 

Sporocleptes Arnold, 1948, Occ. Pap. natn. Mus. Sth Rhod. 2 (14): 219. Type-species: Sporo- 
cleptes nicotiana Arnold, 1948, loc. cit., by original designation. 

Crateropsis Patrizi, 1948, Boll. 1st. Ent. Univ. Bologna 17 : 174 [as a subgenus of Solen- 
opsis]. Type-species: Solenopsis (Crateropsis) elementeitae Patrizi, 1948, loc. cit., by original 
designation. Provisional synonymy of Ettershank, 1966, Aust. J. Zool. 14 : 120. 

Nimbamyrma Bernard, 1952, Mfon. Inst, franc. Afr. noire 19 : 240. Type-species: Nimbamyrma 
villiersi Bernard, 1952, op. cit. : 241, by monotypy. Provisional synonymy of Ettershank, 
1966, Aust. J. Zool. 14 : 120. 

Median portion of clypeus longitudinally bicarinate. Palp formula 2,2 in major workers 
(Ettershank, 1966). Antennae 9-, 10- or n- segmented with a club of two segments. Eyes 
present, small. Promesonotal suture absent from dorsum of alitrunk; metanotal groove 
impressed. Propodeum armed with a pair of teeth or at least sharply angulate. Dimorphic 
species without intermediates, the major workers with massive heads. 

Ants of this genus are small to minute and usually nest in rotten wood to which 
the bark is still adherent. Workers are quite common in Berlese samples of leaf 
litter and have been found in epiphytic moss on trees. 

PAEDALGUS Forel 

Paedalgus Forel, 1911, in Escherich, Termitenleben auf Ceylon : 217. Jena. Type-species: 
Paedalgus escherichi Forel, 1911, op. cit. : 218, by monotypy. 

Mandibles with four teeth. Palp formula 2,2, (Ettershank, 1966). Clypeus longitudinally 
bicarinate. Eyes present, minute, of two ommatidia only. Antennae 9-segmented with 
club of two segments. Promesonotal suture absent; metanotal groove not impressed. Dorsum 
of alitrunk with sharp lateral margins; propodeum unarmed. 

Minute yellowish ants nesting in the walls of termitaria or rotten wood infested 
by termites. 

PHEIDOLOGETON Mayr 

Pheidologeton Mayr, 1862, Verh. zool.-bot. Ges. Wien 12 : 750. Type-species: Oecodoma diversa 
Jerdon, 1851, Madras J. Lit. & Sci. 17 : 109, by designation of Bingham, 1903. 



354 B. BOLTON 

Amauromyrmex Wheeler, 1929, Am. Mus. Novit. no. 349 : 1. Type-species: Amauyomyrmex 
speculifrons Wheeler, 1929, loc. cit. [= Pheidolo^eton silenns (F. Smith, 1858)], by original 
designation. 

Idrisella Santschi, 1937, Annls Soc. ent. Belg. 77 : 372. Type-species: Pheidologeton dentiviris 
Forel, 1913, Arch. Naturgesch. 79 : 192, by original designation. 

Polymorphic. Mandibles of major workers often edentate or with reduced, rounded teeth; 
minor workers with five or six teeth. Palp formula 2,2 (Ettershank, 1966). Clypeus not 
bicarinate. Eyes present. Antennae 11 -segmented with a 2-segmented club. Promesonotal 
suture absent in minor workers, becoming increasingly developed with increased worker size. 
Metanotal groove impressed; propodeum bispinose. 

Nests in and under rotten logs ; uncommon in West Africa. 

SOLENOPSIS Westwood 

Solenopsis Westwood, 1841, Ann. Mag. nat. Hist. 6 : 87. Type-species: Solenopsis mandibularis 
Westwood, 1841 [= Atta geminata Fabricius, 1804, Syst. Piez. : 243], by monotypy. 

Diplorhoptrum Mayr, 1855, Verh. zool.-bot. Ges. Wien 5 : 449. Type-species: Formica fugax 
Latreille, [1798], Essai . . . Fourmis de la France : 46, Brive, by monotypy. 

Synsolenopsis Forel, 1918, Bull. Soc. vaud. Sci. nat. 52 : 155 [as a subgenus of Solenopsis]. 
Type-species: Solenopsis bruchiella Emery, 1921, Genera Insect. Myrmicinae, fasc. 174A: 
199, nom. substit., pro Solenopsis bruchi Forel, 1918 (nee Solenopsis bruchi Forel, 1912), 
by monotypy. 

Diagyne Santschi, 1923, Revue Suisse Zool. 30 : 267 [as a subgenus of Solenopsis]. Type- 
species: Solenopsis succinea Emery, 1890, Boll. Soc. ent. Ital. 22 : 52, by original designation. 

Labauchena Santschi, 1930, Revta Soc. ent. argent. 13 : 81. Type-species: Labauchena daguerrei 
Santschi, 1930, loc. cit., by monotypy. 

Euophthalma Creighton, 1930, Proc. Am. Acad. Arts Sci. 66 : 43 [as a subgenus of Solenopsis]. 
Type-species: Myrmica globularia F. Smith, 1858, Cat. Hym. Brit. Mus. 6 : 131, by original 
designation. 

Oedaleocerus Creighton, 1930, Proc. Am. Acad. Arts Sci. 66 : 43 [as a subgenus of Solenopsis]. 
Type-species: Solenopsis angulata Emery, 1894, in von Ihering, Berl. ent. Z. 39 : 393, by 
original designation. 

Bisolenopsis Kusnezov, 1953, Acta Zool. lilloana 13 : 1. Type-species Bisolenopsis sea Kusnezov, 
1953, loc. cit., by monotypy. 

Paranamyrma Kusnezov, 1954, Mems Mus. Entre Rios 30 : 9. Type-species: Paranamyrma 
solenopsidis Kusnezov, 1954, °P- cr t- : 12, by monotypy. 

Lilidris Kusnezov, 1957, Zool.Anz. 158 : 268, 274. Type-species : Lilidris metatarsalis Kusnezov, 
1957. hoc. cit., by monotypy. 

Granisolenopsis Kusnezov, 1957, Zool. Anz. 158 : 270, 277 [as a subgenus of Solenopsis]. Type- 
species: Solenopsis (Granisolenopsis) granivora Kusnezov, 1957, op. cit. : 278, by monotypy. 

Monomorphic or polymorphic. Mandibles with three or four teeth. Palp formula 1,2, the 
maxillary palp geniculate (Ettershank, 1966). Clypeus strongly longitudinally bicarinate, the 
median area sharply elevated and deeply inserted between the frontal carinae. Antennae 
10-segmented with a 2-segmented club. Promesonotal suture absent from dorsum of ali- 
trunk; metanotal groove impressed; propodeum unarmed. 

The genus is represented by only three or four indigenous species in West Africa, 
which are small yellowish ants nesting in soil at the bases of trees or in the leaf- 
litter. The species are general scavengers and are often found on dead insects 
lying in the leaf-litter at the bases of trees. One species, S. geminata (Fabricius) has 
been introduced to West Africa from the neotropical region. It is very common 
in the Warri Delta in Nigeria, where it is called 'okubrass'. 



ANT GENERA OF WEST AFRICA 355 

SYLLOPHOPSIS Santschi 

Syllophopsis Santschi, 1915, Annls Soc. ent. Fr. 84 : 259 [as a subgenus of Monomoriiim]. Type- 
species: Monomorium modestum Santschi, 1914, Meddn Goteborgs Mus. Zool. Afd. 3 : 17, 
by monotypy. 

Syllophopsis Santschi; Santschi, 1921, Annls Soc. ent. Belg. 61 : 120. [Raised to genus.] 

As Monomorium but median portion of clypeus somewhat swollen, without longitudinal 
carinae. Antennae always 12-segmented with a 3-segmented club. 

Tribe TETRAMORIINI 

DECAMORIUM Forel 

Decamorium Forel, 1913, Annls Soc. ent. Belg. 62 : 121 [as a subgenus of Tetramorium]. Type- 
species: Tetvamorium {Decamorium) decern Forel, 1913, loc. cit., by monotypy. 
Decamorium Forel; Wheeler, 1922^, Bull. Am. Mus. nat. Hist. 45 : 906. [Raised to genus.] 

Mandibles with five to six teeth; palp formula 4,3. Sides of median portion of clypeus and 
posterior margins of the lateral portions raised, bordering the antennal insertions. Antennae 
10-segmented with a 3-segmented club. Antennal scrobes present above the eyes, the ventral 
margins of the scrobes very poorly defined. Anterodorsal pronotal angles acute; promesonotal 
suture absent from dorsum of alitrunk. Metanotal groove impressed; propodeum armed with 
a pair of teeth, metanotal lobes present. Sting with a triangular lamelliform appendage 
apicodorsally. Femora of legs swollen. 

The single species D. decern (Forel) nests in rotten logs or twigs buried in the leaf- 
litter. Foragers often leave the nest and progress through the leaf-litter in single 
file. The species has been observed preying on termites. 

RHOPTROMYRMEX Mayr 

Rhoptromyrmex Mayr, 1901, Annln naturh. Mus. Wien 16 : 18. Type-species: Rhoptromyrmex 
globulinodis Mayr, 1901, op. cit. : 20, by designation of Wheeler, 191 1. 

Acidomyrmex Emery, 1915, Bull. Soc. ent. Fr. 1915 : 191 [as a subgenus of Rhoptromyrmex']. 
Type-species: Rhoptromyrmex wroughtonii Forel, 1902, Revue Suisse. Zool. 10 : 231, by original 
designation. 

Palp formula 3,2 (Brown, 1964). Posterior margins of lateral portions of clypeus raised into 
ridges in front of antennal insertions. Antennae twelve-segmented with a three-segmented 
club. Frontal carinae short, divergent behind; antennal scrobes absent. Sutures absent on 
dorsum of alitrunk, but metanotal groove usually visible. Propodeum unarmed in African 
species. 

Collections of Rhoptromyrmex species in West Africa are uncommon but occasion- 
ally occur in leaf-litter samples, usually from the forest zone. 

TETRAMORIUM Mayr 

Tetramorium Mayr, 1855, Verh. zool.-bot. Ges. Wien 5 : 423. Type-species: Formica caespitum 
Linnaeus, 1758, Syst. Nat., ed. 10 : 581, by designation of Girard, 1879. 

Tetrogmus Roger, Berl. ent. Z. 1 : 10. Type-species: Tetrogmus caldarius Roger, 1857, op. cit.: 
12 [= Myrmica simillima F. Smith, 1851], by monotypy. 

Mandibles with three or four large teeth apically, followed by a variable number of denticulae. 
Palp formula 4,3. Posterior margins of lateral portions of clypeus raised into a ridge bordering 



356 B. BOLTON 

the antennal insertions. Antennae 12-segmented with a 3-segmented club. Antennal scrobes 
absent to feebly present; the frontal carinae often extended backwards as a pair of diverging 
rugae which may run to the occipital margin of the head. Anterolateral angles of the pro- 
notum characteristically sharply angulate, giving a square-shouldered appearance in dorsal 
view. Sutures absent from dorsum of alitrunk, although the metanotal groove is usually 
impressed. Propodeum armed with a pair of spines or teeth, metapleural lobes present, acute, 
may project as a pair of spines below those of the propodeum. Sting with an apicodorsal 
triangular lamelliform appendage. Femora of legs usually distinctly swollen. Setal develop- 
ment not as in Triglyphothrix below. 

Common ants varying from medium to small in size. Usually nesting in rotten 
wood or in compressed leaf-litter, but a few species nest directly into open ground, 
some are arboreal and some are termitolestic. The T. sericeiventre Emery group of 
species includes elongate, long-legged ants making crater nests in open ground. 
They are mostly confined to savannah but also occur in forest clearings and on 
paths. T. termitobium Emery and allies are small species usually found nesting in 
rotten logs with termites, or in the walls of termitaria. 



TRIGLYPHOTHRIX Forel 

Triglyphothrix Forel, 1890, Annls Soc. ent. Belg. 34 : 106. Type-species: Triglyphothrix walshi 
Forel, 1890, op. cit. : 107, by monotypy. 

As Tetramorium above but antennal scrobes usually well developed, deep and divided into 
upper and lower compartments by a longitudinal ridge. Setae trifid or multifid, best observed 
by viewing the alitrunk and pedicel in profile at magnifications of 80 x or more. Hairs usually 
abundant, giving the ant a furry appearance to the naked eye. 

In a few species the antennal scrobes are poorly developed but in these the trifid hair character 
is apparent. 

Leaf-litter and ground foraging species nesting directly into the soil or in rotten 
wood. 



XIPHOMYRMEX Forel 

Xiphomyrmex Forel, 1887, Mitt, schweiz. ent. Ges. 7 : 385 [as a subgenus of Tetramorium]. 
Type-species: Tetramorium (Xiphomyrmex) kelleri Forel, 1889, loc. cit., by designation of 
Wheeler, 191 1. 

Xiphomyrmex Forel; Wheeler, 1922, Bull. Am. Mus. nat. Hist. 45 : 906. [Raised to genus.] 

As Tetramorium above but antennae 11 -segmented with a 3-segmented club. Antennal 
scrobes usually better developed, at least with a distinct dorsal boundary. Palp formula 
usually 4,3 .rarely reduced to 3,3. Sting with a spatulate appendage apically, projecting from 
the body of the sting at a shallow angle. 

A number of species completely lack sculpture and are highly polished, usually jet-black, 
but most species have the head and alitrunk with some sculpturation. In some species one or 
both segments of the pedicel may be squamiform, this character usually corresponding to loss of 
sculpture. 

Nests are made in rotten wood and foragers are quite common in the leaf -litter 
and in log-mould. 



ANT GENERA OF WEST AFRICA 357 

Subfamily DORYLINAE 

Tribe AENICTINI 

AENICTUS Shuckard 

Aenictus Shuckard, 1840, Ann. nat. Hist. 5 : 266. Type-species: Aenictus ambiguus Shuckard, 

1840, loc. cit., by original designation. 
Typhlatta F. Smith, 1857, /. Proc. Linn. Soc. 2 : 79. Type-species: Typhlatta laeviceps F. 

Smith, 1857, l° c - c it-, by monotypy. 
Paraenictus Wheeler, 1929, Boll. Lab. Zool. gen. agr. Portici 24 : 27 [as a subgenus of Aenictus]. 

Type-species: Aenictus (Paraenictus) silvestrii Wheeler, 1929, op. cit. : 28, by monotypy. 

Palp formula 2,2 (Gotwald, 1969). Posterior margin of clypeus and lobes of frontal carinae 
raised, forming a ridge around the antennal insertions, which are exposed in dorsal view. Gena 
laterad of each antennal insertion with a carina of variable length. Antennae ten-segmented. 
Eyes absent. Dorsum of alitrunk without sutures, metanotal groove usually impressed. Pedi- 
cel of two segments, the petiole usually sessile in front. Pygidium not impressed, without 
teeth or spines. 

Small to medium sized ants, yellowish or reddish brown in colour, often with large 
areas of cuticle unsculptured, especially on the head, pronotum and gaster. The 
ants follow an 'army ant' lifeway and their narrow marching columns are quite 
common in the leaf-litter layer. They may also be found in and under rotten logs, 
under bark, and amongst the roots of trees and bushes. 

Tribe DORYLINI 
D ORYL US Fabricius 

Dorylus Fabricius, 1793, Ent. Syst. 2 : 365. Type-species: Vespa helvola Linnaeus, 1764, 

Mus. Ludov. Ulr. : 412, by designation of Shuckard, 1840, Ann. nat. Hist. 5 : 315. 
Sphegomyrmex Imhoff, 1852, Verh. naturf. Ges. Basel 10 : 175. Type-species: Dorylus nigricans 

Illiger, 1802, Magazin Insectenk. 1 : 188, by monotypy. 
Cosmaecetes Spinola, 1853, Memorie Accad. Sci. Torino (2) 13 : 70. Type-species: Cosmaecetes 

homalinus Spinola, 1853, op. cit. : 71 [= Typhlopone fulva Westwood 1840, Introd. Class. 

Insects 2 : 219], by monotypy. 
Shuckardia Emery, 1895, Zool. Jb. Abt. Syst. 8 : 703, 740. Type-species: Alaopone abeilli 

E. Andre, 1885 [= Dorylus atriceps Shuckard, 1840, Ann. nat. Hist. 5 : 323], by original 

designation. 

Polymorphic. Mandibles with apical tooth long and acute, with at least one other tooth on 
the inner margin, usually with more. Larger workers have fewer teeth than smaller forms. 
Palp formula 2,2 (Gotwald, 1969). Frontal carinae vertical, the condylar bulbs of the antennae 
exposed. Antennae 9-, 10- or 11-segmented; eyes absent. Genae not longitudinally carinate. 
Promesonotal suture present, mobile; metanotal groove absent. Pedicel of a single segment, 
the first gastral segment somewhat reduced, smaller than the second segment. Pygidium 
impressed, armed at each side with a tooth or spine. 

The larger species, placed in the subgenus Anomma Shuckard, are the well known 
Driver Ants, their trails often being seen crossing paths where covered runways 
are built by the ants to shield them from direct sunlight. 

Numerous species in other subgenera are never seen on the surface of the ground 
by day. They are found in rotten logs, tree-stumps, leaf -litter and termitaria and 
occasionally under bark or in the earth beneath stones and logs. All species are 
carnivorous and indulge in nomadism and group predation. 



358 B. BOLTON 

Subfamily LEPTANILLINAE 

Tribe LEPTANILLINI 

LEPTANILLA Emery 

Leptanilla Emery, 1870, Boll. Soc. ent. Hal. 2 : 196. Type-species: Leptanilla revelierii Emery, 
1870, loc. cit., by monotypy (nee Leptanilla Holmgren, 1908). 

Frontal carinae raised so that the condylar bulbs of the antennae are exposed in dorsal 
view. Antennae 12-segmented. Eyes absent; genae not carinate. Promesonotal suture 
present, metanotal groove absent. Pedicel of two segments. Pygidium not impressed nor 
armed with spines or teeth laterally. 

Minute ants usually with a total length of less than 1 -5 mm. Depigmented, colour yellow. 

These very small ants carry out an army ant lifeway in the soil and are probably 
predacious on the interstitial fauna of the soil. At present known only from Ghana 
where they were recovered from a Berlese funnel sample taken from a cocoa farm 
in the eastern region of that country. 

Subfamily PSEUDOMYRMECINAE 

Tribe PSEUDOMYRMECINI 

P ACHY SIM A Emery 

Pachysima Emery, 1912, Annls Soc. ent. Belg. 61 : 97 [as a subgenus of Sima]. Type-species: 

Tetraponera aethiops F. Smith, 1877, Trans, ent. Soc. Lond. : 71, by original designation. 
Pachysima Emery; Donisthorpe, 1916, Entomologist's Rec. J. Var. 28 : 242. [Raised to genus.] 

Mandibles with four or five teeth; palp formula 5,4. Antennae 12-segmented without a 
differentiated club. Eyes well developed occupying about one third of the length of the side 
of the head and situated behind the midlength of the head. Three well developed ocelli present. 
Promesonotum distinct, metanotum present as a sclerite on the dorsum of the alitrunk. Middle 
and hind legs with one large pectinate and one small simple spur; claws armed with a tooth 
close to the apex. Petiole and postpetiole each with acute ventral processes. 

Black ants, medium to large in size (total length 7 to 14 mm), living usually in 
plants of the genus Barteria, and tending large cocci ds on the plant. 

TETRAPONERA F. Smith 

Tetraponera F. Smith, 1852, Ann. Mag. nat. Hist. (2) 9 : 44. Type-species: Tetraponera atrata 
F. Smith, 1852, loc. cit. [= Eciton nigrum Jerdon, 1851, Madras J. Lit. Soc. 17 : 111], by 
designation of Wheeler, 191 1. 

Sima Roger, 1863, Berl. ent. Z. 7 : 178. Type-species: Sima compressa Roger, 1863, op. cit. : 179 
[= Pseudomyrma? allaborans Walker, i860], by monotypy. 

As Pachysima above but slender, more elongate ants with proportionally shorter legs. Clypeus 
sometimes produced into a spine or armed with a row of teeth or with a crenulate anterior margin. 
Eyes larger than in Pachysima, often occupying one half the length of the side of the head. 
Ocelli variously developed, often absent but there may be one, two or three present. Either 
the petiole alone with a ventral process or both segments of the pedicel without ventral processes. 

Arboreal species nesting in hollow twigs and branches, their colonies extending 
deep into the trunk if hollow or rotten. Foragers sometimes descend to ground 
level where they may be found close to the base of the trunk or on surface roots. 
The ants are very active with rapid jerky movements and abrupt changes of direction. 



ANT GENERA OF WEST AFRICA 359 

VITICICOLA Wheeler 

Viticicola Wheeler, 1920, Psyche, Camb. 27 : 53. Type-species: Sima tessmanni Stitz, 1910 
Mitt. zool. Mus. Berlin 5 : 131, by original designation. 

Palp formula 3,3 (Wheeler, 1922). Antennae 12-segmented with a 3-segmented club. Eyes 
small, occupying less than one third of the side of the head and situated at about the midlength 
of the head. Anterior ocellus present or with all ocelli absent. Claws simple. Otherwise as 
Pachysima above, but decidely smaller, total length less than 7 mm. 

Living only in the hollow stems of the plant Vitex staudtii Guerke. 

Subfamily DOLICHODERINAE 

Tribe TAPINOMINI 

ENGRAMMA Forel 

Engramma Forel, 1905, Annls Soc. ent. Belg. 49 : 180. Type-species: Engramma lujae Forel, 
1905, op. cit. : 181, by monotypy. 

Mandibles with apical and subapical teeth large, followed by a series of denticulae. Anterior 
border of median portion of clypeus deeply and strongly emarginate. Palp formula 4,3. Eyes 
well developed, set at or just in front of the midlength of the head and on the dorsal surface. 
Antennae 12-segmented. Promesonotal and metanotal sutures present on dorsum of alitrunk. 
Petiole reduced and overhung by the first gastral segment dorsally, invisible in dorsal view. 
Gaster with five segments visible in dorsal view, anal and associated orifices apical. 

Wheeler (19226 : 202) states that most species live in the cavities of myrmeco- 
phytes but that one lives in the ground and another inhabits a woven nest mixed with 
vegetable fibres, attached to the trunks of trees. 

IRIDOMYRMEX Mayr 

Iridomyrmex Mayr, 1862, Verh. zool.-bot. Ges. Wien 12 : 702. Type-species: Formica detecta 
F. Smith, 1858, Cat. Hym. Brit. Mus. 6 : 36 [= Formica purpurea F. Smith, 1858], by desig- 
nation of Bingham, 1903. 

Doleromyrma Forel, 1907, Annls hist.-nat. Mus. natn. hung. 5 : 28 [as a subgenus of Tapinoma]. 
Type-species: Tapinoma (Doleromyrma) darwinianum Forel, 1907, loc. cit., by monotypy. 

Mandibles with the two apical teeth enlarged, the remainder of the apical margin with a 
series of denticulae of varying sizes. Anterior clypeal margin shallowly concave. Palp 
formula 6,4. Eyes located on the dorsal surface of the anterior half of the head. Antennae 
12-segmented. Petiole a small but distinct scale, inclined forewards but not overhung by the 
first gastral segment. 

The single species of this genus represented on the African continent is /. humilis 
(Mayr), an introduction from the neotropical region. To the present time the 
species has not been successfully introduced to West Africa, although it is well 
established in Southern Africa. 

TAPINOMA Forster 

Tapinoma Forster, 1850, Hym. Stud. 1 : 43. Aachen. Type-species: Tapinoma collina 
Forster, 1850 [= Formica erratica Latreille, 1798, Hist. Nat. Form. : 182], by monotypy. 

Micromyrma Dufour, 1857, Annls Soc. ent. Fr. 5 : 60. Type-species: Tapinoma dufouri Donis- 
thorpe, 1943, Ann. Mag. nal. Hist. (11) 10 : 662. 



360 B. BOLTON 

Mandibles with apical two or three teeth large, followed by a row of denticles. Palp formula 
6,4. Clypeus with or without median anterior border emarginate. Antennae 12-segmented. 
Eyes placed at or in front of the midlength of the side of the head on the dorsal surface. Pro- 
podeum unarmed or rarely with a pair of blunt tubercles. Petiole reduced or vestigial, over- 
hung by the first gastral segment and not visible in dorsal view. Gaster in dorsal view with 
four visible tergites; anal and associated orifices ventral. 

Nests under bark, in rotten wood, in compressed leaf-litter or in the soil. T. 
melanocephalum (F.) is a common species in houses in West Africa. 

TECHNOMYRMEX Mayr 

Technomyrmex Mayr, 1872, Ann. Mus. civ. Stor. nat. Giacomo Doria 2 : 147. Type-species: 
Technomyrmex strenua Mayr, 1872, loc. cit., by designation of Bingham, 1903. 

Aphantolepis Wheeler, 1930, Psyche, Camb. 37 : 44. Type-species: Aphantolepis quadricolor 
Wheeler, 1930, loc. cit., by original designation. 

As Tapinoma but with five gastral segments visible in dorsal view, the anal and associated 
orifices apical. 



Subfamily FORMICINAE 

Tribe CAMPONOTINI 

CAMPO NOT US Mayr 

Camponotus Mayr, 1861, Enrop. Fovmicid. : 35. Wien. Type-species: Formica ligniperda 
Latreille, 1802, Fourmis : 88, by designation of Bingham, 1903. 

Polymorphic. Mandibles with five to seven stout teeth. Clypeus often with median portion 
projecting as a truncated lobe, more obvious in larger workers. Palp formula 6,4. Antennae 
12-segmented, without clubs, inserted some distance behind the posterior clypeal margin 
(a distance usually greater than the basal width of the scape). Frontal carinae converging 
anteriorly, leaving the antennal insertions only partially or not at all covered. Eyes present, 
usually well developed, situated behind the midlength of the head. Promesonotal suture 
distinct, development of other sutures variable. The mesoscutellum is often present on the 
dorsal surface of the alitrunk in larger workers and in some the metanotum is also present on 
the dorsum. Alitrunk unarmed, without spines or teeth on pronotum or propodeum, although 
the latter may be abruptly truncated. Petiole a node or scale, without spines or teeth. Acido- 
pore circular, clearly visible, not concealed by the pygidium. 

Common; nests are formed in rotten wood, in the earth, in rotten branches and 
twigs of standing trees, or directly into living wood by extending the galleries begun 
by wood-boring beetles. Medium to very large ants (5 to 15 mm) found in all 
localities. A number of species are arboreal whilst others are purely nocturnal in 
habits, but the majority are diurnal and terrestrial. All are active, fast-running 
ants with powerful mandibles. 



PHASMOMYRMEX Stitz 

Phasmomyrmex Stitz, 1910, Mitt. zool. Mus. Berlin 5 : 146. Type-species: Phasmomyrmex 
sericeus Stitz, 1910, loc. cit. [= Camponotus buchneri Forel, 1886, Annls Soc. ent. Belg. 30 : 
183], by monotypy. 



ANT GENERA OF WEST AFRICA 361 

Monomorphic. Mandibles with five teeth. Clypeus with median portion broadly and 
shallowly excised. Palp formula 6,4. Antennae 12-segmented, without clubs. Insertions of 
antennae, form of frontal carinae and eyes as in Camponotus. Anterodorsal pronotal angles 
projecting as short teeth or with the sides strongly marginate. Metanotal groove impressed, 
the propodeum truncated posteriorly, unarmed. Petiole a node extended into a short dorso- 
lateral tooth on each side. Acidopore circular, not concealed by the pygidium. 

Arboreal, nesting and foraging in trees, very rarely descending to ground level. 

POLYRHACHIS F. Smith 

Polyrhachis F. Smith, 1857, /■ Proc. Linn. Soc. 2 : 58. Type-species: Formica bihamata Drury, 

1773, Illust. Nat. Hist. vol. 2 : 73, by original designation. 
Hoplomyrmus Gerstaecker, 1858, Mbev. dt. Akad. Wiss. Berl. : 262. Type-species: Hoplo- 

myrmus schistacea Gerstaecker, 1858, loc. cit., by monotypy. 

Mandibles usually with five teeth, rarely with four. Median portion of clypeus usually pro- 
jecting as a lobe, rarely broadly emarginate. Palp formula 6,4. Antennae 12-segmented, 
not clubbed, inserted some distance behind the posterior clypeal margin (a distance usually 
greater than the basal width of the scape). Eyes well developed, often strongly protuberant, 
situated on the posterior half of the head. Anterolateral pronotal angles projecting as teeth 
or spines. Alitrunk most often marginate laterally; the propodeum usually bispinose or 
bidentate (very rarely otherwise). Petiole armed with from two to six teeth or spines. Acido- 
pore concealed by the pygidium when not in use. 

Arboreal, usually nesting in the wood or in rot holes in the trunk or branches. 
Some species build nests of silk mixed with vegetable fibres adherent to the under- 
sides of leaves. A few species nest in the earth. Black, agile ants which may have 
strikingly coloured, dense pubescence (gold or silver) on the alitrunk or gaster. 

Tribe FORMICINI 
CAT A GLYPHIS Forster 

Cataglyphis Forster, 1850, Verh. naturh. Ver. preuss. Rheinl. 7 : 493. Type-species: Formica 
megacola Forster, 1850, op. cit. : 490. [= Cataglyphis fairmairei Forster, 1850, loc. cit.], 
by monotypy. 

Mandibles strongly dentate; palp formula 6,4. Antennae 12-segmented, inserted very close to 
the posterior clypeal margin (a distance less than the basal width of the scape) . Eyes and ocelli 
present, the eyes situated behind the midlength of the head. Petiole a node. Acidopore 
borne on a conical projection of the hypopygium and surrounded by a fringe of hair. Legs very 
long. 

Large rust-red ants confined to savannah and semi-desert conditions; occurring 
on the coastal plains. Crater nests are made directly into the earth. The ants are 
very active and fast running. 

Tribe LASIINI 
PARATRECHINA Motschulsky 

Paratrechina Motschulsky, 1863, Bull. Soc. Nat. Moscou 36 (3) : 13. Type-species: Paratrechina 
currens Motschulsky, 1863, op. cit. : 14 [= Formica longicomis Latreille, 1802, Fourmis : 
113], by designation of Wheeler, 191 1. 



362 B. BOLTON 

Mandibles narrow, armed with five or six teeth, weakly or not at all overhung by the clypeus. 
Anterior clypeal margin entire or weakly emarginate medially. Palp formula 6,4. Antennae 
12-segmented, inserted close to the posterior margin of the clypeus. Eyes well developed, 
set at or in front of the midlength of the head. Ocelli absent. Propodeum unarmed; petiole 
a reduced scale, inclined forwards and often overhung by the first gastral segment; always 
unarmed above. Acidopore borne on a conical projection of the hypopygium, surrounded by a 
fringe of hairs. Dorsal surface of head, alitrunk and gaster with distinctly paired, coarse 
setae. 

Nests usually in soil or compressed leaf-litter, less frequently in rotten wood or 
twigs. 



PRENOLEPIS Mayr 

Prenolupis Mayr, 1861, Europ. Formicid. : 35. Wien. Type-species: Tapinoma nitens Mayr, 
1852, Verh. zool.-bot. Ver. Wien 2 : 144, by designation of Bingham, 1903. 

As Paratrechina above but the eyes are behind the midlength of the head. The dorsal sur- 
faces of the head, alitrunk and gaster usually have setae but these are never distinctly arranged 
in pairs and are usually fine. 



PSEUDOLASIUS Emery 

Psendolasius Emery, 1877, Ann. Mus. civ. Stor. nat. Giacoma Doria 24 : 244. Type-species: 
Formica familiaris F. Smith, i860, /. Proc. Linn. Soc. 5 : 68, by designation of Bingham, 
1903. 

Polymorphic. Mandibles usually with five or six, rarely with seven or eight teeth, set upon 
an oblique apical border. Palp formula 3,4 in largest workers, 3,3 in smallest. Clypeus well 
developed, overhanging the mandibles. Antennae 12-segmented, their insertions virtually 
confluent with the posterior clypeal margin. Major workers with small eyes present, situated 
at or just in front of the midlength of the head and on the dorsal surface. Minor workers without 
eyes. Petiole a scale which may be inclined forwards. Acidopore borne on a conical projection 
of the hypopygium and surrounded by a fringe of hairs. 

Depigmented, yellowish coloured ants nesting in or under very rotten wood in or 
the soil amongst the roots of trees where dead wood is present. The workers actively 
avoid light but may be found on the surface of the ground during the night. Work- 
ers are often found in Berlese funnel samples of leaf-litter or log-mould. 



Tribe MYRMELACHISTINI 
APHOMOMYRMEX Emery 

Aphomomyrmex Emery, 1899, Annls Soc. ent. Belg. 43 : 493. Type-species: Aphomomyrmex 
afer Emery, 1899, op. cit. : 494, by designation of Wheeler, 191 1. 

Polymorphic. Mandibles with three teeth. Antennae 9-segmented, without a differentiated 
club. Eyes well developed, oval, situated on the dorsal surface of the head at about the mid- 
length. Ocelli present. Pro- and mesonotum in profile strongly convex. Scale of petiole 
high and narrow, emarginate dorsally. 

Arboreal. Small (3-0-3-5 mm) black ants with the legs and antennae paler, reddish. Sides 
of alitrunk smooth and shining, dorsal surfaces of head, alitrunk and gaster finely punctate. 



ANT GENERA OF WEST AFRICA 363 

Tribe OECOPHYLLINI 
OECOPHYLLA F. Smith 

Oecophylla F. Smith, i860, /. Proc. Linn. Soc. 5 : 101. Type-species: Formica smaragdina 
Fabricius, 1775, Syst. Ent. : 828, by monotypy. 

Dimorphic. Mandibles elongate triangular, apical teeth long, acute and crossing over at 
rest. Apical margin behind the first tooth with seven to ten smaller teeth or denticles, of which 
the first and third are usually the largest. (This may not be true of minor workers.) Palp 
formula 5,4. Clypeus large, convex, its anterior margin overhanging the basal borders of the 
mandibles. Antennae 12-segmented, the first funicular segment longer than the second and 
third together. Eyes well developed, ocelli absent but shallow pits may mark their location in 
the major workers. Alitrunk strongly constricted in the mesonotal region, the pronotum 
and propodeum considerably broader than the region separating them. Petiole elongate and 
narrow in dorsal view, forming a low, rounded node in profile. Gaster with acidopore visible, 
not hidden by the pygidium. 

Arboreal ants, making nests by binding leaves together with larval silk. The 
major workers are general carnivores and scavengers, the minors are rarely seen away 
from the nest. The single West African species 0. longinoda (Latreille) tends large 
coccids, often building protective silk tents over large aggregations of coccids. 
The construction of the petiole allows reflexion of the gaster over the alitrunk, a 
position which the ants take when disturbed. 

Tribe PLAGIOLEPIDINI 
ACANTHOLEPIS Mayr 

Acantholepis Mayr, 1861, Europ. Formicid. : 42. Wien. Type-species: Hypoclinea frauenfeldi 
Mayr, 1855, Verh. zool.-bot. Ges. Wien 5 : 378, by monotypy. 

Mandibles with apical margin oblique, dentate, overhung by the clypeus. Palp formula 6,4. 
Antennae 11-segmented. Eyes well developed, ocelli present but may be reduced. Alitrunk 
constricted in the mesonotal region, the propodeum swollen and bidentate or bituberculate. 
Petiole a scale with the dorsal margin bispinose, bidentate or emarginate. Acidopore borne on 
a conical projection of the hypopygium, surrounded by a fringe of hairs. 

Medium sized to small ants, usually black in colour but some species brown or 
yellowish. Nests are made in rotten wood either in standing trees or on the ground, 
or are built directly into hard-packed earth. The foragers of the larger species 
often ascend trees to tend aphids or coccids whilst those of other species are found 
only in the leaf-litter layer. 

ACROPYGA Roger 

Acropyga Roger, 1862, Berl. ent. Z. 6 : 242. Type-species Acropyga acutiventris Roger, 1862, 
op. cit. : 243, by monotypy. 

Mandibles narrowly triangular, with five teeth, not overhung by the clypeus. Palp formula 
2,3 (Wheeler, 1922) or 1,3 (Gotwald, 1969). Antennae 11-segmented. Eyes small, situated in 
front of the midlength of the side of the head; ocelli absent. Alitrunk not constricted in the 
mesonotal region. Propodeum unarmed; petiole an unarmed scale. Acidopore borne on a 
conical projection of the hypopygium surrounded by a fringe of hairs. 

Small, depigmented, yellowish ants. Hypogaeic, uncommon. 



364 B. BOLTON 

PLAGIOLEPIS Mayr 

Plagiolepis Mayr, 1861, Europ. Formicid. : 52. Wien. Type-species: Formica pygmaea 
Latreille, [1798], Essai . . . Fourmis de la France: 45. Brive, by monotypy. 

Mandibles with five teeth; clypeus large and projecting over the basal borders of the man- 
dibles. Palp formula 6,4. Antennae 11-segmented. Eyes well developed, situated in the 
middle of the sides of the head; ocelli usually absent but may be present. Alitrunk short, 
weakly constricted between the pronotum and propodeum. Propodeum unarmed; petiole a 
reduced scale, inclined forwards and may be overhung by the first gastral segment, but never 
armed or emarginate. Acidopore borne on a conical projection, surrounded by a fringe of 
hairs. 

Medium to small ants, monomorphic or polymorphic. Nests are made under the 
bark of trees, in rotten wood or twigs, or in hard-packed earth. 

Tribe SANTSCHIELLINI 
SANTSCHIELLA Forel 

Santschiella Forel, 1916, Revue Suisse Zool. 24 : 434. Type-species: Santschiella kohli Forel, 
1916, op. cit. : 435, by monotypy. 

Mandibles with seven or eight teeth. Antennae 12-segmented. Eyes very large, occupying 
almost the whole of the side of the head; ocelli present. Head bordered posteroventrally by a 
transparent ridge which ends abruptly and is followed by a tooth. Propodeum bispinose. 
Node of petiole truncated posteriorly, armed posterodorsally by an obtuse tooth on each side. 

Rare, believed to be arboreal. 



REFERENCES 

Arnold, G. 1915. A monograph of the Formicidae of South Africa. Ann. S. Afr. Mus. 
14, part 1 : 1-158, 1 pi., 8 figs. 

1916, ibid, part 2 : 159-270, pis 2-4, figs 9-18. 

1917, ibid, part 3 : 271-402, fig. 19. 

1920, ibid, part 4 : 403-578, figs 20-57. 

1922, ibid, part 5 : 579-674, fig. 58. 

1924, ibid, part 6 : 675-766, pis 5-9, fig. 59. 

1952. The genus Terataner Emery. /. ent. Soc. Sth. Afr. 15 : 129-131, 1 fig. 

Bernard, F. 1952. La reserve naturelle integrate du Mt. Nimba. fasc. 1, 11, Hymenoptera, 

Formicidae. MSm. Inst.fr. Afr. noire 19 : 165-270, 3 pis, 15 figs. 
Bingham, C. T. 1903. Fauna of British India, Hymenoptera, vol. 2, Ants and Cuckoo 

Wasps. 506 pp., 1 pi., illustrations. London. 
Brown, W. L., Jr. 1948. A preliminary revision of the higher Dacetini. Trans Am. ent. 

Soc. 74 : 101-129, 2 figs. 
1949a. Revision of the ant tribe Dacetini I. Fauna of Japan, China and Taiwan. 

Mushi 20 : 1-25, 2 figs. 
1949&. Revision of the ant tribe Dacetini III. Epitritus Emery and Quadristruma new 

genus. Trans Am. ent. Soc. 75 : 43-51, 1 fig. 

• 1952a. Revision of the ant genus Serrastruma. Bull. Mus. comp. Zool. Harv. 107 : 65-86. 

1952b. Contributions towards a reclassification of the Formicidae I. Tribe Platythreini. 

Breviora no. 6: 1-6. 
I953 a - Revisionary studies in the ant tribe Dacetini. Am. Midi. Nat. 50 : 1-137, 3 pis. 

10 figs. 



ANT GENERA OF WEST AFRICA 365 

Brown, W. L., 1953k. Characters and synonymies among the genera of ants. Parti. Bveviora 

no. 11 : 1-13. 
J 953 c - Characters and synonymies among the genera of ants. Part II. Bveviora no. 

18 : 1-8. 
1954- The ant genus Strumigenys F. Smith in the Ethiopian and Malagasy Regions. 

Bull. Mits. comp. Zool. Harv. 112 : 1-34, 1 fig. 
1958. Contributions towards a reclassification of the Formicidae II. Tribe Ectatommini. 

Bull. Mus. comp. Zool. Harv. 118 : 173-362, 48 figs. 
i960. Contributions toward a reclassification of the Formicidae III. Tribe Amblyoponini. 

Bull. Mus. comp. Zool. Harv. 122 : 145-230, 48 figs. 
1963. Characters and synonymies among the genera of ants. Part III. Some members 

of the tribe Ponerini. Breviora no. 190 : 1-10. 

1964. Revision of the genus Rhoptromyrmex . Pilot. Reg. Zool. cards 11-19. 7 figs. 

Ithaca, N.Y. 

1971. Characters and synonymies among the genera of ants. Part IV. Some genera of 

the subfamily Myrmicinae. Breviora no. 365 : 1-5. 
Eidmann, H. 1944. Die Ameisenfauna von Fernando Poo. Zool. Jb., Syst. 76 : 413-490, 

2 pis, 17 figs. 
Ettershank, G. 1966. A generic revision of the world Myrmicinae related to Solenopsis 

and Pheidologeton. Aust. J. Zool. 14 : 73-171, 141 figs. 
Gotwald, W. H., Jr. 1969. Comparative morphological studies of the ants with particular 

reference to the mouthparts. Mem. Cornell Univ. Agric. Exp. Stn 408 : 1-150, 374 figs. 
Hung, A. C. F. & Brown, W. L., Jr. 1966. Stmcture of the gastric apex as a subfamily 

character in the Formicinae. Jl N.Y. enl. Soc. 74 : 198-200. 
Menozzi, C. 1929. Revisione delle formiche del genre Mystrium Roger. Zool. Anz. 82 : 

518-536. 9 figs. 
1942. Formiche dell Tsola Fernando Poo e del territorio del Rio Muni (Guinea Spagnola). 

Zool. Anz. 140 : 164-182, 4 figs. (Published 1950). 
Santschi, F. 1914. Formicides de 1'Afrique occidentale et australe. Boll. Lab. Zool. gen. 

agr. Portici 8 : 309-385. 

1924a. Revue du genre Plectroctena F. Smith. Revue Suisse Zool. 31 : 155-173, 3 figs. 

19246. Revision des Myrmicaria d'Afrique. Annls Soc. ent. Belg. 64 : 133-176, 2 figs. 

Taylor, R. W. 1965. A monographic revision of the rare tropicopolitan ant genus Probolo- 

myrmex Mayr. Trans R. ent. Soc. Lond. 117 : 345-365, 35 figs. 
1967. A monographic revision of the ant genus Ponera Latreille. Pacif. Insects Monogr. 

13 : 1-112, 87 figs. 
Weber, N. A. 1943. Ants of the Imatong Mountains, Anglo-Egyptian Sudan. Bull. Mus. 

comp. Zool. Harv. 93 : 261-389, 16 pis. 
1950. African species of the genus Oligomyrmex Mayr. Am. Mus. Novit. no. 1442 : 1-19, 

20 figs. 

1952. Studies on African Myrmicinae. Am. Mus. Novit. no. 1548 : 1-32, 36 figs. 

Wheeler, W. M. 191 1. A list of the type species of the genera and subgenera of the Formi- 
cidae. Ann. N.Y. Acad. Sci. 21 : 157-175. 

1922a. Ants of the Belgian Congo. Part I. Distribution of ants of the Ethiopian and 

Malagasy Regions. Bull. Am. Mus. nat. Hist. 45 : 13-37. 

1922b. Ants of the Belgian Congo. Part II. Ants collected by the American Museum 

Congo Expedition, ibid. : 39-269, 23 pis, 76 figs. 

1922c. Ants of the Belgian Congo. Part VII. Keys to the genera and subgenera of 

ants. ibid. : 631-710. 

19220'. Ants of the Belgian Congo. Part VIII. Synonymic list of the ants of the 

Ethiopian Region, ibid. : 711-1004. 

Wilson, E. O. & Taylor, R. W. 1967. Ants of Polynesia. Pacif. Insects Monogr. 14 : 
1-109, 84 figs. 



3 66 



B. BOLTON 



INDEX 



Acantholepis, 363 
Acidomyrmex, 355 
Acrocoelia, 342 
Acropyga, 363 
Adelomyrmex, 346 
Aenictus, 357 
Aeromyrma, 352 
Aethiopopone, 341 
Amauromyrmex , 354 
Amblyopone, 330 
Aneleus, 353 
Anochetus, 333 
Aphantolepis, 360 
Aphomomyrmex, 362 
Arotropus, 330 
Asphinctopone, 334 
Atopomyrmex, 348 

Bisolenopsis, 354 
Bondroitia, 351 
Bothroponera, 334 
Brachyponera, 335 

Cacopone, 335 
Calyptomyrmex, 347 
Camponotus, 360 
Cardiocondyla, 341 
Carebara, 351 
Cataglyphis, 361 
Cataulacus, 342 
Caulomyrma, 346 
Centromyrmex, 335 
Cephaloxys, 345 
Cerapachys, 340 
Codiomyrmex, 343 
Corynomyrmex, 352 
Cosmaecetes, 357 
Crateropsis, 353 
Cratomyrmex, 349 
Crematogaster, 342 
Cryptocephalus, 348 
Cryptopone, 336 

Decamorium, 355 
Diagyne, 354 
Diplomorium, 351 
Diplorhoptrum, 354 
Discothyrea, 332 
Dodous, 348 
Doleromyrma, 359 



Names printed in italics are recognised synonyms. 

Dorylozelus, 337 
Dorylus, 357 



Emeryia, 341 
Eneria, 345 
Engramma, 359 
Epipheidole, 350 
Epitritus, 343 
Epixenus, 352 
Epoecus, 352 
Equestrimessor , 352 
Erebomyrma, 353 
Ericapelta, 330 
Escherichia, 332 
Euophthalma, 354 

Fulakora, 330 

Glyphopone, 336 
Goniothorax, 346 
Granisolenopsis, 354 

Hendecatella, 353 
Heptacondylus, 349 
Hercynia, 350 
Holcomyrmex, 352 
Hoplomyrmus, 361 
Hylidris, 348 
Hypoponera, 336 

Idrisella, 354 
Ireneidris, 352 
Iridomyrmex, 359 
I solcomyrmex , 352 

Labauchena, 354 
Labidogenys, 345 
Lampromyrmex, 352 
Lecanomyrma, 353 
Lepidopone, 334 
Leptanilla, 358 
Leptogenys, 337 
Leptomyrma, 350 
Leptopone, 336 
Leptothorax, 346 
Lilidris, 354 
Limnomyrmex , 346 
Lithomyrmex, 330 



INDEX 



367 



Macromischoides, 347 

Megaloponera, 337 
Megaponera, 337 
Melissotarsus, 347 
Meranoplus, 348 
Mesoponera, 338 
Messor, 350 
Miccostruma, 343 
Microbolbos, 337 
Microdaceton, 344 
Micromyrma, $y> 
Monomorium, 351 
Myrmapatetes, 333 
Myrmicaria, 349 
Mystrium, 331 



Promyopias, 339 
Proscopomyrmex , 345 

Protamblyopone 330 
Psalidomyrmex, 339 
Pseudolasius, 362 
Pseudoneoponera , 334 
Pseudosphincta, 332 
Pseudosysphincta, 332 
Pyramica, 345 



Quadristruma, 344 



Khoptromyrmex, 355 



Neoamblyopone, 330 
Nimbamyrma, 353 



Octella, 353 
Odontomachus, 333 
Odontomyrmex , 348 
Oecophylla, 363 
Oedaleocerns, 354 
Oligomyrmex, 352 

Pachysima, 358 
Paedalgus, 353 
Paltothyreus, 338 
Paraenictus, 357 
Paranamyrma, 354 
Paraphacota, 352 
Paratrechina, 361 
Parholcomyrmex , 352 
Pedetes, 333 
Phacota, 351 
Phasmomyrmex, 360 
Pheidole, 350 
Pheidologeton, 353 
Phrynoponera, 338 
Phyracaces, 340 
Physatta, 349 
Plagiolepis, 364 
Platythyrea, 331 
Plectroctena, 339 
Polyrhachis, 361 
Prenolepis, 362 
Pristomyrmex, 348 
Probolomyrmex, 332 
Proceratium, 333 
Prodiscothyrea, 332 



Santschiella, 364 
Serrastruma, 344 
Shuckardia, 357 

Sima, 358 
Simopone, 341 
Smithistruma, 345 
Solenops, 353 
Solenopsis, 354 
Spalacomyrmex . 335 
Spelaeomyrmex , 353 
Sphegomyrmex , 357 
Sphinctomyrmex, 341 
Sporocleptes, 353 
Stigmatomma, 330 
Strumigenys, 345 
Syllophopsis, 355 
Synsolenopsis, 354 
Sysphincta, 333 
Sysphingta, 333 



Tapinoma, 359 

Technomyrmex, 360 
Terataner, 349 
Tetramorium, 355 
Tetraponera, 358 
Tetrogmus, 355 
Trachymesopus, 340 
Tranopeltoides, 342 
Trichomyrmex, 351 
Trichoscapa, 345 
Triglyphothrix, 356 
Typhlatta, 357 
Typhloteras, 335 



Viticicola, 359 



368 INDEX 

Wasmannia, 350 Xenhyboma, 352 

Weberidris, 347 Xeromyrmex, 352 

Wheeleria, 352 Xiphomyrmex, 356 

Wheeleriella, 352 Xvninier, 330 



B. Bolton, B.Sc, A.R.C.S. 

Department of Entomology 

British Museum (Natural History) 

Cromwell Road, London SW7 5BD 



A LIST OF SUPPLEMENTS 

TO THE ENTOMOLOGICAL SERIES 

OF THE BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 
18 plates, 270 text-figures. August, 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 : 500 text-figures. September, 

1965- £3-25- 

5. Ahmad, I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156: 
475 text-figures. November, 1965. (out of print) £2.15. 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. 

£3-i5. 

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 
text-figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera) . Pp. 509. £8.50. Reprinted 1972. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322: 348 text-figures. August, 1967. £8. 

11. Mound, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172: 
82 text-figures. May, 1968. £4. 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. 
November, 1968. £5. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
Nymphalidae) . Pp. 155: 3 plates, 101 text-figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North-Western Europe 
(Hymenoptera : Chalcidoidea) . Pp. 908: 686 text-figures. November, 1969. 

£i9- 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 

68 plates, 15 text-figures. October, 1971. £12. 

18. Sands, W. A. The Soldierless Termites of Africa (Isoptera Termitidae). 
Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. 



Jfljjj Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 



12 I 4 w 



- 

CONTRIBUTIONS TOWARDS 



1 2 i c:c, 



REVISION OF MYRSIDEA 

WATERSTON. VII. (PHTHIRAPTERA: 

AMBLYCERA: MENOPONIDAE) 



i 



B. K. TANDAN 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 27 No. 7 

LONDON : 1972 



CONTRIBUTIONS TOWARDS A REVISION OF 

MYRSIDEA WATERSTON. VII. (PHTHIRAPTERA: 

AMBLYCERA: MENOPONIDAE) 



BY 



BHUP KISHORE TANDAN 

University of Lucknow 



Pp 369-410; 2 Plates, 54 Text-figures 




BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 27 No. 7 

LONDON : 1972 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 27, No. 7 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Ent.). 



Trustees of the British Museum (Natural History), 1972 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 18 December, 1972 Price £1.95 



CONTRIBUTIONS TOWARDS A REVISION OF 

MYRSIDEA WATERSTON. VII. (PHTHIRAPTERA 

AMBLYCERA: MENOPONIDAE) 

By B. K. TANDAN 



CONTENTS 



Synopsis 
Introduction 
Taxonomic Characters 
Species Descriptions 



Key to the Species of Myrsidea Parasitic on Garrulax and Pomatorhinas 401 

403 
404 
406 
406 
407 
410 



Host-parasite List- 
Discussion 
Acknowledgments 
References . 
Tables 
Index 



Page 
37i 
37i 
37i 
374 



_ SYNOPSIS 

This part of a series of papers on Myrsidea Waterston deals with the species parasitic on bab- 
blers of the genera Gamilax and Pomatorhinus (subfamily Timaliinae, Aves). It includes a 
redescription of one known and descriptions of 12 new species, a key to the species, a host- 
parasite list and a short discussion on host-parasite relationships. 

INTRODUCTION 

This is part VII in the series of papers initiated by Clay (1966). Like the previous 
part (Tandan & Clay, 1971) this too is devoted to species of Myrsidea parasitic on 
babblers (subfamily Timaliinae of Muscicapidae) and deals with species parasitizing 
the genera Garrulax and Pomatorhinus. Although this part was intended exclusively 
for species parasitizing Garrulax, species from Pomatorhinus have been included as 
they were found to resemble closely those occurring on certain species of Garrulax. 
The host-names are according to Deignan in the Check-List of Birds of the World 
(1964), and the names of Provinces of Thailand are also as given in Deignan (1963). 
The following abbreviations have been used for the collections which are the deposi- 
tories for the material on which this study is based: BMNH, British Museum (Nat- 
ural History); USNM, U.S. National Museum; EC, K.C. Emerson collection; REC, 
Robert E. Elbel collection. 

TAXONOMIC CHARACTERS 

The characters common to all or most of the species dealt with in this paper are 
given below. Neither these characters nor the generic characters given by Clay 
(1966 : 330-332 and 1969) are repeated in the specific descriptions. 



372 B. K. TANDAN 

1. Head of the same general shape (PI. I, figs 55-57), differences shown by measure- 
ments. Head chaetotaxy basically as in M. thoracica (see Clay, 1966, fig. 1), as also 
the relative proportions of the pair of setae on the last segment of the maxillary palp 
but in some taxa there may be greater individual variation. End of seta 10 usually 
reaches to about the middle of seta 11 (Text-fig. 10). Gular setae usually 4 + 4, 
occasionally 4 + 3 or 4 -\- 5 and exceptionally 3 + 3, 5 + 5 or 64-5. Antenna as 
in Clay, 1966, fig. 2, and the two sensilla coeloconica (Clay, 1969 : 8, 1970 : 76) on 
the second segment slightly apart. Hypopharynx variable. 

2. Thorax. Pronotum with 3 + 3 setae near each antero-lateral corner of which, 
according to available evidence, the 2 + 2 outer spiniform ones may be propleural 
setae and the relatively 1 + 1 inner moderately long to long pronotal setae. Pos- 
terior margin of pronotum with 3 + 3 long and stout setae. Mesonotum undivided. 
Metanotum normal or modified with 3 + 3 antero-lateral spiniform and 1 + 1 long 
postero-lateral setae (not included in setal counts), lying between the latter are a 
variable number of marginal setae. Metapleural setae spiniform. Metasternal 
plate triangular in shape with a varying number of setae, its anterior width depending 
on that of the metanotum but the posterior narrow apex variable in length. One or 
two of the metasternal setae each side are anterior and more central than the re- 
maining somewhat marginal ones; only in one species was the number of anterior 
setae over 4 (Text-figs 14, 15). First tibia with 3 + 3 outer ventro-lateral and a 
varying number of dorso-lateral setae. 

3. Abdomen. In both sexes spiracles open on the tergites; in the female either IV 
or V, in the male V, is the broadest tergite. In the female the anterior terga are 
normal or modified, the extent of modification being reflected by the curvature of the 
line of marginal setae; the edge of the vulva may be smooth or serrated. Micro- 
trichia of the inner surface of the genital chamber variable in the three species in 
which they are visible. No structure resembling the complicated bursa copulatrix 
of forms infesting Turdoides (Timaliinae) and Icteridae (Clay, 1968 ; Tandan & Clay, 
1 971) surrounds the opening of the spermathecal duct in the genital chamber. The 
duct either appears to open in a depression at the apex of a papilla (Text-fig. 37), 
the genital papilla (sensu Keler, 1971 : 10), with the opening being usually visible, or 
in a thin-walled sac, without the opening being visible. While the genital papilla 
may or may not have characteristic pigmentation, the duct proximal to the opening 
is usually wider and appears to be somewhat hardened. On the other hand, the 
structurally simpler, thin-walled sac is probably the homologue of the bursa copul- 
atrix as indicated by the presence of concretions in the sac of one species (orientalis) 
resembling those seen inside the bursa of diverse species parasitic on Turdoides. 
Owing to the membranous nature of the duct and sac (or bursa copulatrix), they are 
not always apparent in all specimens, so that in those species (patkaiensis and 
macraidoia) in which one or both these structures have not been seen, they may in 
fact be present. However, the details of these structures differ in different individuals 
of the same species (Text-figs 33, 34), due mainly to distortion produced during pre- 
paration of the specimen, and as the number of specimens is small, these limitations 
prevent a satisfactory comparison between populations from different hosts. 

The male genitalia have all the basic components composing the external genitalia 



REVISION OF MYRSIDEA 373 

of Myrsidea. These show more specific variation than has been found in recent 
studies of groups of species from related hosts, in which it is usually only the genital 
sclerite which shows specific differences. A long and narrow spermatophore closely 
associated with the genital sclerite has been seen in some species. 

Abdominal chaetotaxy. Tergites without anterior setae. Post-spiracular setae 
III and V always shorter and finer than II and IV, III being slightly longer than V. 
In the male on tergum IX 1 + 1 (occasionally 3) moderately long to long marginal 
and 4 + 4, exceptionally 9 or 10, short internal anal setae. Anterior setae absent or 
present on pleurites II-VII, or some of them. Pleurite VIII has 3 + 3 setae, the 
central one being very long and stout, but the lengths of the outer and inner ones 
relative to each other vary; extra inner setae (v) are usually present in the female, 
rarely in the male (Text-fig. 27). Sternite I without setae. Arrangement of setae 
on sternites IV-VII in the female, IV-VIII in the male and the genital region in both 
sexes as in Myrsidea from Trudoides (see Tandan & Clay, 1971), only the setae in the 
genital region being much stouter. The outermost setae on the vulval margin 
stouter than the inner ones. 

The following abdominal setae are found in all the species dealt with here :i + i 
antero-lateral, spiniform on tergite I; I + I post-spiraculars on I-VIII and the as- 
sociated spiniform of the post-spiracular setal complex (Clay, 1970) on II-VIII; in 
the male usually 1 -f- 1, occasionally 1 + 2, moderately long to long on posterior 
margin of IX ; 2 + 2 antero-lateral, spiniform or short on sternite II ; 1 -f- 1 postero- 
lateral on sternite VIII in the male and 2 + 2 in the genital region of both sexes; 
3 + 3 laterals on segment IX, the 2 + 2 outer and relatively dorsal ones being 
probably pleural. These setae have been excluded from the setal counts given in the 
text, Key and Tables, with the exception of the 1 + 1 postero-lateral setae on sternite 
VIII in the male and 2 + 2 in the genital region, which may not be readily identifiable 
when there are numerous setae in this part. The 2 + 2 setae on sternite II, usually 
separable from other setae by their proportions and antero-lateral position, may not 
always be separable on one or both sides when the number of setae on this sternite 
is greater (Text-fig. 29). 

A combination of some of the following taxonomic specific characters has been 
found useful in determining the status of populations from different hosts, the 
relatively more important ones having been marked with an asterisk (*). 1. Degree 
of development of hypopharynx*. 2. Form of anterior terga and nature of the 
vulval margin in the female*. 3. Form of metanotum in the female and shape of 
metasternal plate. 4. Shape of components of male genitalia, especially the genital 
sclerite*. 5. Details of the structure associated with the opening of the sper- 
mathecal duct*. 6. Number of dorso-lateral setae on tibia I and in the brush on 
femur III. 7. Number of setae on metanotum and metasternum. 8. Length of 
setae on pleurite I in the female ; presence or absence of anterior setae on pleurites 
II-VII and the relative proportions of the outer and inner setae on pleurite VIII. 
9. Number of setae on tergum IX in the female, on I and of terminal setae on IX in 
the male. 10. Number and length of setae on tergum I in the female and on VIII 
in both sexes*. 11. Length of post-spiracular setae III, V and VI. 12. Number 
of setae on sternite II in both sexes and also III in the female*. 13. Arrangement 



374 B. K. TANDAN 

of setae on sternites II and III in the female*. 14. Presence or absence of 
central anterior setae on sternites III or IV-VI. 15. Number of setae in the genital 
region and on vulval margin*. 



SPECIES DESCRIPTIONS 

The following species-descriptions and measurements (in millimetres, usually 
corrected to two decimal places) are on the same lines as those given by Clay (1966, 
1968) and the Tables of the sternal chaetotaxy, based on specimens from the type- 
host only, by Tandan & Clay (1971). Figures in parentheses denote the number of 
specimens or structures examined or measured, and x the mean. The chaetotaxy 
in the figures agrees with the specimens from which these were drawn and broken or 
missing setae, shown by broken lines, have been usually completed from the other 
side of the same specimen or from another specimen. 



Myrsidea sehri Ansari, 1951 
(PI. 2, fig. 59; Text-figs 10, 21, 39) 

Type-host : Garrulax I. lineatus (Vigors) . 

Myrsidea sehri Ansari, 195 1 : 177, fig. 19. Holotype $, from Trochalopteron lineatum grisescentior 
(Hartert), India (BMNH) [examined]. 

Only the two specimens comprising the type-material of this species, the first 
Myrsidea to be described from one of the Timaliinae, have been available for study. 
These are in poor condition with most of the thoracic and abdominal setae missing 
and it has not been possible to take all the necessary measurements or to determine 
the size o